sign in sign up
<<
Home , Bahia (plant), Encelia, Geraea, Geraea canescens, Iostephane, Phoebanthus

Systematics of (Compositae-) Author(s): David M. Spooner Source: Systematic Botany Monographs, Vol. 30, Systematics of Simsia (Compositae- Heliantheae) (Nov. 12, 1990), pp. 1-90 Published by: American Society of Taxonomists Stable URL: http://www.jstor.org/stable/25027790 Accessed: 08/07/2010 11:04

Your use of the JSTOR archive indicates your acceptance of JSTOR's Terms and Conditions of Use, available at http://www.jstor.org/page/info/about/policies/terms.jsp. JSTOR's Terms and Conditions of Use provides, in part, that unless you have obtained prior permission, you may not download an entire issue of a journal or multiple copies of articles, and you may use content in the JSTOR archive only for your personal, non-commercial use.

Please contact the publisher regarding any further use of this work. Publisher contact information may be obtained at http://www.jstor.org/action/showPublisher?publisherCode=aspt.

Each copy of any part of a JSTOR transmission must contain the same copyright notice that appears on the screen or printed page of such transmission.

JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected].

American Society of Plant Taxonomists is collaborating with JSTOR to digitize, preserve and extend access to Systematic Botany Monographs.

http://www.jstor.org SYSTEMATICS OF SIMSIA (COMPOSITAE-HELIANTHEAE)

David M. Spooner Vegetable Crops Research Unit Agricultural Research Service, U.S.D.A. Department of Horticulture University ofWisconsin Madison, Wisconsin 53706

ABSTRACT.Simsia (Compositae-Heliantheae) comprises annuals, herbaceous perennials, suffru tescent perennials, and . It occurs throughout tropical America from sea level to 3000 m. This study is based on comparative morphology, including extensive field studies and observations in the greenhouse, phenetic studies, crossing experiments, and cytology. Eighteen species including seven varieties are recognized. Two species, S. santarosensis and S. villasenorii, are newly described. The following new combinations are proposed: S. annectens var. grayi, S. foetida var. grandiflora, S. foetida var. jamaicensis, S. foetida var. megacephala, and S. foetida var. panamensis. Twenty species and varieties are predominantly self-incompatible and outcrossers; three species are self-compatible (or apomictic) and possibly inbreeders.Meiosis is normal inmost of the F1 hybrids produced from outcross ing species but irregular in F1 hybrids involving a self-compatible species. Chromosome numbers are uniformly n = 17. Counts are reported for the first time for the following: S. annectens var. annectens, S. chaseae, S. foetida var. panamensis, S. holwayi, S. molinae, S. santarosensis, S. setosa, S. tenuis, and S. villasenorii.

INTRODUCTION

Simsia is a of herbs and shrubs distributed from the southwestern to the center of . The greatest diversity occurs from central to Panama, where 18 of the 23 species and varieties grow. Some of the species (e.g., S. amplexicaulis, S. lagascaeformis) arewidespread roadside and agriculturalweeds and are familiar to collectors of Mexican Compositae. Others (e.g., S. holwayi, S. steyermarkii) are narrowly restricted endemics and are seldom encoun tered. Many of the taxa occur in dry, tropical deciduous forests (e.g., S. foetida), others in upland pine-oak to cloud forests (e.g., S. amplexicaulis); S. eurylepis occurs along the humid Mexican Gulf coastal lowlands. The species share a 2-4 seriate involucre, linear, sterile ray florets (except for one eradiate species, S. eurylepis), a convex receptacle with conduplicate, medium-textured, persistent pales, fertile disc florets with a well-defined, glandular-puberulent tube and abruptly expanded throat, and laterally flattened, ovate to obovate to elliptic disc achenes with or without awns (rarely with intermediate squamellae). The genus was last monographed by Blake (1913) who resurrected it from synonymy within Adanson and recognized 22 species and three varieties. In a recent useful and insightful synopsis of Mexican and Central American species, Robinson and Brettell (1972) treated 24 species from these areas, including seven new ones, and estimated 35 species total for the genus. This synopsis was based on limited material (only US), did not include field-investigations, and did not treat species from and . Despite previous work on Simsia, basic taxonomic problems remain, as also

1 2 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 30

indicated by Nash (1976) and by Jansen and Stuessy (1980). The genus is taxonomi cally difficult by any standard. The polymorphic nature of many species has led to a confusing array of names and taxa of uncertain circumscription.A possible contrib uting factor to this taxonomic confusion is the apparent tendency of certain species to hybridize, a pattern discerned from herbarium investigations and further sug gested by field- and greenhouse-studies. The present study of Simsia is based on considerably more data than available to previous workers. Over 3600 herbarium specimens have been examined from 63 institutions. In addition, Kunth and related genera have been examined at ENCB, GH, MEXU, MO, OS, and US for a study of generic relationships. Five field tripswere conducted over a total of eight months from 1982-1990 throughout Mexico, , , , , Panama, , Ecua dor, and Argentina; these resulted in field-observations and collections of all taxa except for one endemic variety from Jamaica. The field-investigations were espe cially useful for observations on intra- and interpopulational variability, habitats, natural hybridization, and for the collection of pertinent morphological observa tions difficult to obtain from herbarium collections, such as habit, plant height, branching patterns, corolla color, and odor. Cytological collections provided material for 95 chromosome counts covering all taxa except S. foetida var. jamaicensis. Prior to this study, 56 chromosome counts were documented for 14 of the 23 species and varieties and one natural hybrid of Simsia; 93 new counts for 22 of these taxa and two natural hybrids (Appendix 1) are listed here. First counts are reported for S. annectens var. annectens, S. chaseae, S. foetida var. panamensis, S. holwayi, S. molinae, S. santarosensis, S. setosa, S. tenuis, S. villasenorii, and the natural hybrids S. am plexicaulis x S. lagascaeformis and S. foetida var. foetida x S. lagascaeformis. It is evident that Simsia has evolved at the diploid or diploidized level, and therefore speciation within the genus must have involved factors other than changes in ploidy level. Numerical phenetic studies have been used to help define taxonomic limits and to develop hypotheses on evolutionary patterns in the genus. Sixteen species and three additional varieties have been grown in the greenhouse; of these, 11 species and three varieties were used in an artificial crossing program. These crossing studies provided data on compatibility systems, genomic compatibility among vari ous species, stability of morphological features, and insights into the nature of taxonomic problems and evolutionary processes operative in Simsia. Simsia is here treated as comprising 18 species, including seven varieties. Two species, S. santarosensis and S. villasenorii, are newly described. The species are all monobasic (x = 17) and homoploid (2n = 34). Three species are self-compatible (or apomictic); all others are self-incompatible. Local hybridization is believed to occur between certain species. One species, S. chaseae, exhibits additive morphological features between S. foetida and S. eurylepsis that could have resulted either by hybrid speciation or by convergent evolution.

TAXONOMIC HISTORY

The taxonomic history of Simsia begins with the publication of Coreopsis foetida Cavanilles (1791), described from garden-grown plants raised from achenes 1990 SIMSIA 3

supplied by the Sesse andMocifio expedition. Persoon (1807) circumscribedSimsia, including S. foetida (Cavanilles) Persoon, and S. heterophylla (Cavanilles) Persoon [= heterophylla (Cavanilles) Bentham ex Hemsley]. Kunth (1818) and Sprengel (1826) submerged Simsia intoXimenesia Cavanilles (= Verbesina L.), but Cassini (1829) recognized Simsia and greatly improved the characterization of the genus. Simsia was accepted as a genus by later workers, and new taxa were described by de Candolle (1836), Gray (1850, 1853), Bentham (1853), and Triana (1858). Gray (1873) later reduced Simsia to a section of Encelia. Subsequently, taxa now assigned to Simsia were described inEncelia (Hemsley 1881;Gray 1887;Rose 1895; Fernald 1897; Kuntze 1898;Hieronymus 1901;Greenman 1903;Millspaugh 1904). A modern interpretation of Encelia and related genera (or genera at least superfi cially similar to Encelia) awaited the insightfulwork of Blake (1913), who resur rected Simsia and described eight new taxa. Simsia has been accepted since that time, with later additions provided by Blake (1917; 1918a; 1924; 1928), Loesener (1923), Cuatrecasas (1954), and Robinson and Brettell (1972).

GENERIC RELATIONSHIPS

Simsia is relatively homogeneous, and, except for some close relatives in Viguiera, it rarely is confused with other genera. The most diagnostic and readily observable characters are the herbaceous, 2-4-seriate, imbricate involucres, the linear ray ovaries, and the laterally flattened disc achenes almost always lacking squamellae. The combination of Simsia and Encelia by Gray (1873) and laterworkers was based on a superficial resemblance of the radiate, paleaceous heads and laterally flattened achenes of these species. Encelia (sensu Blake, 1913) differs by its alter nate and pales deciduous with the villous, white-margined achenes; also, the base chromosome number of species of Encelia is x = 18. Encelia is now recognized to be part of an assemblage that includes A. Nelson, Flourensia de Candolle, Torrey & Gray, and, perhaps, Phoebanthus S. F. Blake (Robin son 1981; Dillon 1984; Clark & Sanders 1986; Sanders & Clark 1987). In addition to the above features, Encelia and Enceliopsis share similar benzofuran chemistry (Mitsakos & Proksch 1985), and Encelia and Geraea sometimes hybridize in nature (Kyhos 1967). More recent hypotheses have linked Simsia to Viguiera (Robinson & Brettell 1972), L. (Robinson 1979), some members Viguiera ser. Grammato glossae S. F. Blake (this study), and Desfontaines ex Gmelin (Schilling & Jansen 1989). Simsia, Helianthus, Tithonia, and much of Viguiera share many of the morphological features that help to characterize Simsia. In addition, these four genera are united by the presence of anthochlors, unknown in the rest of the Heliantheae except for Coreopsidinae (Crawford& Stuessy 1981). Differences among Tithonia, Simsia, Helianthus, and some species of Viguiera ser. Grammatoglossae are few. Tithonia is distinguished by a fistulose . The most diagnostic characters of the latter three groups are in the disc achenes. Those of Simsia are laterally very flattened, thickened only in the central area, with semi-persistent awns present or absent, and only rarely with shorter squamellae between the awns. The achenes of Helianthus and Viguiera, although laterally 4 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 30 compressed, are plump and biconvex throughout. Viguiera usually has persistent awns and additional, shorter squamellae, and Helianthus has caducous awns, usu ally without squamellae (Heiser et al. 1969). Additional differences among these genera are less diagnostic, but they also help delimitation. Simsia has long (3-10.2 mm), linear ray ovaries, whereas those of Helianthus and much of Viguiera are shorter or on the low end of this range and often more obovate. The style branches of Simsia are long (2-4 mm), thin, and grade to an attenuate apex with a poorly defined appendage formed by papillae at the end. Those of Helianthus andmuch of Viguiera are generally shorter and more deltoid, and inHelianthus often have a well-defined appendage. In addition to these features, many specimens of Simsia possess characteristic foliaceous nodal discs (see section Morphology and Taxo nomic Criteria), unknown inHelianthus and rare inViguiera. My investigations suggest that Simsia ismost closely related to some members of Viguiera ser. Grammatoglossae. Blake (1918b) characterized this series by its herbaceous, lanceolate to oblong phyllaries, which contrastwith themore indurate phyllaries of other taxa. He included 11 species of herbs, suffrutescent perennials, and shrubs, distributed from to Guatemala. Most species within this series, unlike other species in Viguiera, have similar elongate style branches, linear ray ovaries, and herbaceous phyllaries resembling those of Simsia. All have biconvex achenes, and none has the nodal discs so characteristic of many species of Simsia. Viguiera ser. Grammatoglossae is probably an unnatural taxon. To this group belongs V. potosina S. F. Blake, placed by Blake (1913) in series Viguiera (=ser. Dentatae S. F. Blake). Viguiera potosina has floral flavonoids (Rieseberg & Schil ling 1985), a chromosome number (n = 17, vs. n = 18 for V. ser. Viguiera; Schilling & Schilling 1986), and herbaceous phyllaries more similar to those of ser. Grammatoglossae and Simsia. In addition, some species appear misplaced in ser. Grammatoglossae. Viguiera parkinsonii (Hemsley) S. F. Blake, V. pringlei Robin son & Greenman, and V. seemannii Schultz-Bipontinus have indurate phyllaries and an overall appearance quite different from other members of this series, and thus are best excluded from it. Another species of ser. Grammatoglossae, V. mima S. F. Blake, known only from the type, bears a superficial resemblance to S. foetida var. grandiflora, but its heads, rays, and achenes resemble those of Tithonia. For these reasons, Simsia is considered closely related to a subset of Viguiera ser. Grammatoglossae, including V. potosina, but excluding V. mima, V. parkin sonii, V. pringlei, and V. seemannii. All species within this subset sharewith Simsia a diploid chromosome number, 2n = 34; V. cordifolia A. Gray (Heiser & Smith 1955; Keil & Stuessy 1977; this study, see Appendix 1), V. grammatoglossa de Candolle (Turner et al. 1962; Ralston et al. 1989), V. ovata (A. Gray) S. F. Blake (Sundberg et al. 1986), and V. potosina (Schilling & Schilling 1986; Ralston et al. 1989). The only differences (although intergrading)between Simsia and these very similar species in Viguiera ser. Grammatoglossae are the lateral thickening and minor shape differences of the achenes (flattened and obovate to elliptic in Simsia, biconvex and obovate to linear-obovate in Viguiera ser. Grammatoglossae), squa mellae of the pappus (usually absent in Simsia, usually present in Viguiera), and the nodal discs (often present in Simsia, rare in Viguiera). The differences between Simsia and Viguiera ser. Grammatoglossae are few and often intergrading, and future study may demonstrate the congeneric status of Simsia and some of these 1990 SIMSIA 5

taxa inViguiera. Other genera similarly are distinguished from Viguiera by few and sometimes intergrading characters, and the problem of intergradinggeneric traits and the problematic placement of taxa extends beyond the Simsia-Viguiera relation ship (Cronquist 1985).

INTERSPECIFIC RELATIONSHIPS

Three distinct phenetic groups are apparent. One group, here informallydesig nated group A, contains S. annectens, S. fruticulosa, S. ghiesbreghtii, S. molinae, S. holwayi, S. setosa, S. tenuis, S. steyermarkii, and S. villasenorii. Most of these species possess a perennial habit, distinct nodal discs, unequal phyllaries (S. annectens var. annectens has subequal phyllaries), and black anthers. Group B contains S. amplexicaulis, S. chaseae, S. dombeyana, and S. foetida. These species share an annual habit, lack nodal discs, and have subequal phyllaries (some populations of S. chaseae have unequal phyllaries), and yellow anthers. The possible hybrid origin of S. chaseae (see section Artificial Hybridizations) makes its placement in Group B problematic, but it has an overall appearance similar to other species inGroup B. Group C contains S. eurylepis and S. lagascaeformis. These species share the annual habit, lack nodal discs, and possess the yellow anthers of group B, but have the unequal phyllaries of group A. The placement of Group C with members of Group A or Group B is problematic because of thismixed distribution of charac ters, but inmy subjective opinion, Group C shows most similaritywith Group B. Simsia lagascaeformis additionally shares with S. amplexicaulis stems that com monly root at the lower nodes and a purple distal band on the anthers. and S. sanguinea are distinctive and are hard to ally with any other species or groups, because both have various combinations of features of Groups A and B, but I intuitively group them with Group A. Robinson and Brettell (1972) also conceptualized phenetic groups in Simsia. My group A corresponds to their groups A and H, and my group B to their group G. They, too, were unable to ally S. calva and S. sanguinea with other groups. Their groups C, D, and F differ from mine. I initially attempted a cladistic analysis. I agree with the goals and approaches of most cladistic schools in attempting to reconstruct phylogeny based on explicit approaches involving the clear presentation of the characters and character states, the polarization of these states by outgroup comparison (e.g., Watrous & Wheeler 1981;Maddison et al. 1984), and the construction of cladograms with repeatable techniques based on parsimony. Among problems encountered in this analysis, the most serious was the presence of polymorphic characters in Simsia. This problem of polymorphic character states is common in flowering plants, and character state coding in these situations has been handled in different ways. Cantino (1982) con verted such qualitative polymorphic characters into meristic ones reflecting the proportion of different character states under consideration. Jansen (1985) coded the most common character state or discarded those that showed much variability. If characters with moderate to high variability in Simsia were discarded, very few characters remained. Another problem was polarization of character states, be cause the outgroup, a subset of Viguiera ser. Grammatoglossae, had few invariant 6 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 30 characters useful for polarizing those in Simsia. Similar problems with character state polarity in the related genus Helianthus led Schilling and Heiser (1981) to analyze the ingroup relationships of Helianthus by an unrooted network. Cladistic analyses were conducted using PAUP version 2.4.1 (Swofford 1984) after coding the characters by theirmost common state, using unrooted characters, and rooting the cladogram with the subset of Viguiera ser. Grammatoglossae men tioned above. The topologies of the resulting cladograms had numerous unresolved branches (polycotomies), and homoplasy (parallelisms and reversals) was high (>70%). The large degree of homoplasy revealed in these analysesmay be real and reflect mosaic evolution (sensu Stebbins 1974) in an evolutionarily active genus. Alternatively, or additionally, this homoplasy may be caused by reticulate evolu tion or introgression that would confound efforts at phylogenetic reconstruction (Wagner 1983; Funk 1985). Similar problems with polymorphic characters with poorly defined character states is present in other groups in the Compositae, e.g., in Senecio L. (Barkley 1988). In addition, minor changes in the data-set (attempted with some of the polymorphic characters) sometimes caused significant differences in the topologies. Because of the problems of utilizing morphology for cladistic analyses, I in tended to present an intuitive classification, recognizing two sections inSimsia, one section composed of members of Group A, S. calva, and S. sanguinea, and another section composed of members of groups B and C. Within the second section, I intended to recognize two series, one series containingmembers of group B and the other members of group C (see Schilling & Spooner 1988). Recent (Schilling & Jansen 1989) and ongoing chloroplast-DNA studieswith Simsia, Viguiera, and other close genera (Schilling, pers. comm.) are providing additional data about phylo genetic relationships inSimsia and related genera. I believe it ispremature to recog nize formally these sections and series in Simsia before these data are available.

NUMERICAL

Numerical taxonomic studies were undertaken to help clarify taxonomic struc ture in the wide-ranging and taxonomically difficult S. foetida complex, to investi gate morphological intermediacy of S. chaseae to S. eurylepis and S. foetida, and to help distinguish S. chaseae from S. dombeyana.

SIMSIA FOETIDACOMPLEX The S. foetida complex is a group of morphologically similar and intergrading taxa that traditionally has been difficult to circumscribe and identify.As recognized here, the S. foetida complex iswell-defined and easily distinguished by a number of shared features. All components are annuals or short-lived, facultative perennials. The stems, leaves; and phyllaries are densely covered with subsessile to stipulate biseriate glands that impart a viscid feel and a range of lemony and spicy odors in living plants. The phyllaries are linear, subequal, often with caudate tips and sinuate margins. The plants occur in hot, lowland habitats, most frequently in areas of tropical deciduous forests, from sea level to ca. 1700 m (occasionally to 2150 m), from , Mexico, to central Panama, with disjunct populations in Jamaica, , and . 1990 SIMSIA 7

Table 1. Characters and states used in numerical phenetics of the Simsia foetida complex, S. chaseae hybridization, and the S. chaseae-S. dombeyana study. Leaves. 1. Blade length (mm). 2. Blade width (mm). 3. length (cm). 4. Interpetiolar stipule height (mm). 5. Glandular pubescence: subsessile (0), stipitate (1). 6. Abaxial non-glandular pubescence: strigose, hispid, or hirsute (0), pilose (1). Peduncles and Heads. 7. Longest peduncle length (cm). 8 Head diameter (mm). 9. Head height (mm). Phyllaries. 10. Number. 11. Outer series length (mm). 12. Outer serieswidth (mm). 13. Inner series width (mm). 14. Ratio inner/outer series lengths. Ray florets. 15. Number. 16. Corolla length (mm). 17. Corolla width (mm). 18. Ovary length (mm). Disc florets. 19. Number. 20. Corolla length (mm). 21. Tube length (mm). 22. Lobe length (mm). 23. Anther length (mm). 24. Achene length (mm). 25. Achene width (mm). 26. Longest awn length (mm). Pales. 27. Outermost pale length (mm). 28. Outermost pale width (mm).

Within this complex, taxonomically significant variation ismainly quantitative. Previous workers, however, have treated this quantitative variation inmore qualita tive or inmore discrete, quantitative ways. Despite considerable intergradation, geographically correlated modes of variation may be recognized. Preliminary stud ies suggested that six taxa could be recognized within this group. Principal compo nents analysis (PCA) was used to gain an overall estimate of these patterns of variation, and stepwise discriminant analysis was then employed to test the signifi cance of the groups. Materials and Methods. One hundred and ten individuals (operational taxo nomic units, OTU's) were measured for 26 quantitative and 2 qualitative morpho logical characters (Table 1). The populations were selected to represent the range of morphological variation and geographical distribution of the group. In six cases, two plants from a population were measured. The measured populations are marked with an asterisk in the list of representative specimens in the taxonomic section. Measurements of head parts were made from collections preserved in FAA or from herbarium material revived in AerosolT (a commercial wetting agent). Heads were chosen from selected collections in a random manner; width and diame ter were from the widest point of a structure. Leaf and nodal disc measurements were taken from the largest structures available. The data were analyzed by PCA available in the NT-SYS package (Rohlf et al. 1972), and a stepwise discriminant analysis, available in the BMDP computer pack age (Dixon et al. 1985). For PCA, the data-matrices were standardized by using the STAND subroutine of NT-SYS, resulting in a mean of zero and a standard devia tion near unity. The PCA was then performed using the subroutine FACTOR. For stepwise discriminant analysis, only the 26 quantitative characters were used (i.e., excluding 5 and 6). Character distributions were tested (using the PD2 program in BMDP) for normality by a t-test for skewness and for kurtosis signifi cantly different from 0 (for p < 0.05), using the PD2 program in BMDP. Non normal distributions were normalized by the TRANSFORM functions of BMDP and these new characters were substituted in the data-set. Six distributions could not be normalized and were discarded. A stepwise discriminant analysis was then performed on this reduced and partially transformed data-set using the P7M pro gram of BMDP. Results and Discussion. The results of the PCA are presented in Fig. 1. Factors 1 and 2 account for 43.9% of the total variation. A third factor raised the total to 8 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 30

2

0.70

0 -0.75-1.23 j A'

L3__E______X-1.72 -0.92 -0.68 -0.34 -0.05 0.25 0.54 0.83 1.12 1.41

* Simsia foetida var. foetida (northern populations) o var. foetida (southern populations) e var. jamaicensis * var. megacephala * var. grandiflora * var. panamensis

FIG. 1. First two principal component axes of OTU's of Simsia foetida var. foetida, var. jamaicensis, var. megacephala, var. grandiflora, and var. panamensis.

54.9% but did not change the overall pattern and is not presented. This analysis indicates some overlap between taxa. Despite this overlap, the distinctness of the six taxa I initially recognized (including the northern and the southern populations of S. foetida, see below) are significant at the 0.001 or 0.005 level by a multiple analysis of variance generated by the stepwise discriminant analysis (Table 2). Of the 21 characters utilized in the multiple analysis of variance analysis, seven exhib ited the greatest discriminant utility: phyllary number, nodal disc length, achene length, awn length, ray corolla width, disc corolla tube length. My initial hypothesis suggested that the S. foetida populations from , Mexico, southeast to , Mexico, (northern populations) were distinct from those from , Mexico, to Guatemala (southern populations). All northern populations (except one, see Taxonomy) lack stipules and nodal discs. The major ity of the southern populations possess stipules and nodal discs, but there is consid erable intra-populational variation in this area, with some plants in a population or 1990 SIMSIA 9

Table 2. F-matrix for a stepwise discriminant analysis of the varieties of Simsia foetida (var. foetida partitioned into northern and southern populations, see text). All values are significant at p less than 0.001 (f = 12), or 0.005 (f = 7.20); degrees of freedom = 7.79.

foetida foetida jamaicensis megacephala panamensis (southern (northern populations) populations) foetida 11.29 (northern pop.) jamaicensis 25.27 23.99 megacephala 22.22 30.91 30.44 panamensis 17.28 40.80 39.06 36.64 grandiflora 19.98 38.78 40.86 27.76 12.76

various nodes on individual plants lacking these structures.The PCA analysis (Fig ure 1) demonstrates extreme overlap of the northern and southern populations, but themultiple analysis of variance (Table 2) discriminates them. I have decided not to recognize the northern and southern populations of S. foetida as distinct taxa, because the results of the PCA and the degree of intrapopulationalvariability in the nodal characters do not support such a separation.

SIMSIA CHASEAE, S. EURYLEPIS,AND S. FOETIDA. Simsia chaseae is intermediate for many morphological features between S. eurylepis and S. foetida, which suggested the possibility of its hybrid origin between these two species. A fuller discussion of this hypothesis is presented under Artificial Hybridizations. Materials andMethods. A PCA was conducted on 12 individuals of S. chaseae, 19 individuals of S. eurylepis, 34 individuals of S. foetida var. foetida, and 33 artificial F1 hybrids produced between S. eurylepis and S. foetida var. foetida (indi viduals used in this analysis aremarked with an asterisk in the lists of representative specimens). Twenty-six quantitative morphological characterswere used (Table 1, characters 5 and 6 excluded). Results. The first two components account for 45% of the total variation (the third component raised the total to 57% but did not change the overall pattern and is not presented). The analysis (Fig. 2) indicates a bimodal distribution of charac ters, with some hybrids more similar to one or the other parent. Some of the hybrids, however, are morphologically almost indistinguishable from naturally oc curring S. chaseae. Further discussion of the results are presented under Artificial Hybridizations.

SIMSIA CHASEAEAND S. DOMBEYANA

Simsia chaseae is remarkably similar S. dombeyana. Both are annuals with glan dular leaveswith unwinged petioles andwithout discs, both have light lemon-yellow ligules, and both are self-compatible and have urceolate heads. It is often difficult to distinguish these species, because the differences are minor. Greenhouse-grown S. dombeyana lacks the strong spicy odors characteristic of S. chaseae and S. foetida. Simsia chaseae grows in the northern hemisphere and from June through November; S. dombeyana occurs in the southern hemisphere and flowers from De 10 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 30

2

0.81

0.04

-0.72

- 1 .1 1 ______IJ__ T 1 r I -0.77 -0.57 -0.37 -0.18 0.02 0.22 0.42 0.61 0.80

* Simsia eurylepis * Artificial hybrids: * X * A S. chaseae * S. foetida var. foetida

FIG. 2. First two principal components axes of OTU's of natural populations of Simsia chaseae, S. eurylepis, S. foetida var. foetida and artificial F1 hybrids of S. eurylepis x S. foetida var. foetida.

cember through May. The main character that previously has distinguished these two species has been phyllary gradation. Simsia chaseae originally was described as hav ing unequal phyllaries, and Blake (1913) used this characteristic to separate the species from S. dombeyana, which has subequal phyllaries. Although all specimens of S. dombeyana appear to possess subequal phyllaries, so do some specimens of S. chaseae (e.g., Purpus 11121, Spooner 2859, 2881). These similarities and overlapping characteristics, and my initial difficulty in distinguishing the species, caused me to use PCA to see if two taxa should be recognized, and to use a stepwise discriminant analysis to discover which characters might best discriminate them. Materials and Methods. The methods are the same as those used for analysis of the Simsia foetida complex, but characters 5 and 6 were excluded. Twenty-one OTU's from 12 populations of S. chaseae and 22 OTU's of 19 populations of S. 1990 SIMSIA 11

dombeyana (markedwith an asterisk in representative specimens), representing the morphological and geographical range of the species, were used. Results and Discussion. The results of the PCA are summarized in Figs. 3 and 4. The first two components account for 47.0% of the total variation; the third component raises the total of 59.9%. These results indicate a pattern of partially defined but overlapping taxa. The stepwise discriminant analysis statistically sup ports the recognition of these groups at the 0.001 level. Although phyllary grada tion is the most reliable discriminator of the taxa, it occasionally fails to distinguish the species and additional characters are needed. These characters, arranged in a decreasing order of discriminant utility (see Table 1) are: 14, 16, 26, 13, 22, 12, 16. The similarity of the taxa could be used to support recognition as varieties, but their usual ease of identification and their distinct ranges more strongly suggest specific status.

ARTIFICIAL HYBRIDIZATIONS

Sporadic hybridization is believed to occur naturally between S. amplexicaulis and S. foetida, S. amplexicaulis and S. lagascaeformis, and between S. foetida and S. lagascaeformis. Simsia chaseae possibly could have arisen by hybrid speciation between S. foetida var. foetida and S. eurylepis. Crossing studieswere conducted to provide insights into possible natural hybridization, compatibility systems, and evo lutionary relationshipswithin the genus. Materials andMethods. Forty-eight parental individuals representing 27 popula tions of 11 species, three additional varieties, and two additional populations of naturally occurring S. foetida x S. lagascaeformiswere used in the crossing program in The Ohio State University greenhouses (Appendix 2). In addition, S. dom beyana (Spooner 2907, 2908), S. foetida var. panamensis (Spooner 2903), S. fruticulosa (Spooner 2953, 2957), S. molinae (Spooner 2910), and S. santarosensis (Spooner 2901) were grown and tested for compatibility but not used in the crossing program. The only taxa of Simsia not grown and observed in the greenhouse were S. foetida var. jamaicensis and S. ghiesbreghtii. Heads were bagged before anthesis, and the bags were temporarily removed only for cross-pollinations, which involved gently rubbing the heads of thematernal parent with a detached head of the paternal parent. Five to 20 heads per individual were kept bagged and never cross-pollinated, and at least three of these heads per individual were pollinated with pollen from a different head of the same plant to test for self-incompatibility. As a further test of self-incompatibility, S. calva, S. foetida var. foetida, S. foetida var. megacephala, S. lagascaeformis, and S. setosa, previously indicated by the bagging experiments to be self-incompatible, were iso lated in cages screened to exclude pollinators and crossed 15 or more times with intraplant pollinations without bagging the heads. Only one crossing attempt was made per head because of the fragile nature of some pedicels. Only some florets were receptive during any one crossing attempt. Emasculation was not conducted, because all species used as maternal parents had low rates of selfing (<1%), and most hybrids were apparent by their morphological intermediacy. Pollen fertility of parents and hybrids was estimated by staining pollen in 2% lactophenol in cotton-blue. Pollen was collected from at least two anthers each from 12 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 30

2

1.17 3

0.80

A

0.43 -A A *~ A ae 0.07 -S

-030 ~AA *AA A A A

A -0.67 I.3 I I / lI I l -0.86 -0.63 -0.40 -0.17 0.06 0.29 0.52 0.74 0.97

3

0.68 4 A

0.39 * * 5

570 01 . IIA 5 a I

AA A S --0.15 A A. 0. 0. 0 A &A4AA A A - --0.43 AAA A A

-0.70 A

-0.86 -0.63 -0.40 -0.17 0.06 0.29 0.52 0.74 0.97

A Simsia chaseae * S. dombeyana

S. FIGS. 3, 4. First three principal component axes of OTU's of Simsia chaseae and dombeyana. 1990 SIMSIA 13

49(lC

FIG. 5. Mean percent pollen stainability for intravarietal (within circles), intervarietal (outside circles), and interspecific crosses (on arrows) of Simsia. The number inparentheses is the number of F,'s obtained for each cross. The arrow points to thematernal parent.

at least two different heads of a plant andmixed together on the same slidewith two drops of stain. Observations were made after 24 hours. This procedure was re peated a second time at least one week later, and the higher of the two numberswas taken as representative of the individual. Only fully formed and darkly stained grains were scored as stained. Results and Discussion. The bagged but uncrossed and intraplant-pollinated heads set full complements of achenes over 80% of the time for S. chaseae, S. dombeyana, and S. eurylepis, which indicates that they are self-compatible or apomictic. All other tested species were essentially self-incompatible, but in one or two instances each, 1-3 fruits were formed on the bagged but uncrossed or intraplant-crossed heads of S. foetida var. grandiflora, S. lagascaeformis, and S. sanguinea, which indicates low levels of self-compatibility or possibly apomixis. Flowering F1'swere produced from in ten intraspecificand 22 interspecific combinations (Figure 5, Appendix 2). Hybrids were intermediate inmost floral and vegetative features, but dominance was noted in the glandular and spicy-smelling leaves of all crosses involving S. foetida. Some of these hybrids resulted from the crosses of morphologically quite distinct parents (e.g., S. setosa or S. villasenorii, crossed with S. chaseae, or S. foetida crossed with S. sanguinea). 14 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 30

Table 3. Wilcoxon two-sample test of in tragroup (1 x 1, 2 x 2) vs. intergroup (1 x 2) differ ences in pollen stainability in Simsia. The first num ber is the interspecificmean pollen stainability, the second number is the rank of the mean.

Wilcoxon Two-Sample Test

n2 (n2+ ' C = + n1n2 2 -E

1 x 1, 2 x 2 1 x 2

34- 6 1-1 37- 7 6-2 48- 8 12-3 62 - 10 21 - 4 67 - 11 30 - 5 49 - 9

n2 nl i R=42 E R=24 The parental taxa have the highest levels of mean pollen-stainability with 95% (range 88-100%). The intra- and intervarietalmean pollen-stainabilities were iden tical to each other, 74.4% each, but the ranges varied from 44 to 99% and 61 to 96%, respectively. The pollen-stainability data support my intuitive hypotheses of evolutionary relationships (see Generic Relationships and Interspecific Relation ships). The grouped (1 x 1, 2 x 2) intragroup mean pollen-stainability values are 49.6% (mean ranges 34-67%) vs. the intergroup (1 x 2) mean pollen-stainability values of 19.8% (mean ranges 1-49%). These values, analyzed by the Wilcoxon two-sample test (Sokal & Rohlf 1981) support these groups at the 0.05 level (Table 3). Meiosis of the microsporocytes was examined for one individual sample floret each (except as noted in Table 4) of the intraspecific and interspecific hybrid combi nations (Appendix 2). Frequent occurrence of "sticky" chromosomes made it diffi cult to distinguish true from apparent multivalents, but univalents were evident at metaphase I, and laggards and also bridges and fragments were evident at late anaphase I or telophase I.Meiosis was normal for all of the hybrids except for four combinations that exhibited various degrees of irregularity (Table 4). Cytological observations of themicrosporocytes indicated that these abnormalitieswere recipro cal translocations and paracentric inversions. Of interest is that three of these four crosses involve S. eurylepis and S. chaseae, two of the three self-compatible species (possibly apomictic, but here assumed to be self-compatible) in the genus (the other self-compatible species, S. dombeyana, was not intercrossed in this program). The results suggest that these self-compatible species are also inbreeders not subjected to selection for chromosomal uniformity present in the outbreeders. A concentrated focus of this study was the possible diploid hybrid speciation of S. chaseae, with S. eurylepis and S. foetida var. foetida as putative parents. All three species are lowland annuals; only S. chaseae and S. eurylepis are found on the Gulf coastal lowlands. Simsia chaseae and S. foetida both are glandular with spicy odors 1990 SIMSIA 15

Table 4. Summary of chromosome associations in artificial interspecificF, hybrids atMetaphase I, and at Anaphase I or Telophase I. All other intervarietal and interspecific hybrids, including crosses with naturally occurring S. foetida x S. lagascaeformis, exhibited normal pairing and disjunction at meiosis. One individual from each cross was examined, except as noted below.

Cross Number of cells Univalents at Number of cells Laggards at Percent examined Metaphase I examined Anaphase I or bridgesa mean (range; Telophase I SD) mean (range; SD) VILL x CHASb 39 0.4 (0-4; 0.9) 140 1.3 (0-5; 1.6) 40 SETO x CHAS 79 1.0 (0-6; 1.3) 128 0.4 (0-3; 0.7) 10 FOET x SETO 26 1.7 (0-6; 1.7) 997 0.7 (0-5; 1.0) 16 FOET x EURYc 59 0.2 (1-2; 0.5) 626 0.6 (0-2; 0.3) 1 a. Percent of cells at Anaphase I or Telophase I that exhibited bridges and fragments. b. Pooled data from two hybrid individuals. c. Pooled data from five hybrid individuals.

and radiate heads, and they frequently have been confused in the herbarium. Simsia chaseae and S. eurylepis share lobed leaves, self-compatibility, and urceolate involucres. Simsia chaseae generally is intermediate in phyllary gradation between S. eurylepis and S. foetida var. foetida. Many other species of flowering plants have been hypothesized to have arisen after diploid interspecific hybridization (Raven 1976; Grant 1981). Some of these hypotheses have been based solely on inference from the additive morphology of the putative parents; others have been tested by additivity of secondary chemistry or isozymes, and some have been tested by crosses of the putative parents. The obvious question in all of these scenarios is how natural hybrids formed in this manner maintain their integrity by reproductive isolation from the parents. Various mechanisms have been proposed, such as natural selection in a new environment (e.g., Randolph et al. 1967; Levin 1967), floral isolation (e.g., Straw 1955; Stebbins & Ferlan 1956), or recombinatinal speciation (e.g., Grant 1954). To test the hypothesis of the origin of S. chaseae via diploid hybrid speciation, crosses were made between plants from two populations each of S. eurylepis (al ways serving as the pollen donor) and S. foetida var. foetida. All 33 F1 hybrids were glandular with the spicy odors of S. foetida and S. chaseae but lacked the self compatibility of the latter species. Variability in other characters included such features as phyllary reflexion, ratio of lengths of inner/outer phyllaries, phyllary pubescence, and the presence or absence of ray florets. Some of the F1 hybrids were completely rayless as is the S. eurylepis parent, others had a full complement of rays as in S. foetida, and still others had both rayed and rayless heads on the same plant, with some heads having only 1-4 ray florets. The PCA of these parents and hybrids, along with S. chaseae (see Numerical Taxonomy, Fig. 2), indicated a bimodal distribution of the morphology of the hybrids, with some plants more similar to one or to the other parent. Some of the hybrids, however, were morphologically almost indistinguishable from S. chaseae. Simsia chaseae is largely isolated geographically on the Yucatan Peninsula, but there is some sympatry with S. eurylepis in . Chromosomal sterility barri ers also may effectively isolate S. chaseae from S. eurylepis (inference from data 16 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 30

from Table 4, but data from S. chaseae-S. eurylepis cross not available). Evidence against the hybridization hypothesis is the absence of self-compatibility in all of the F1 hybrids, and the current lack of sympatry of the two putative parents. The hypothesis depends upon the previous co-occurrence of the parents and the attain ment of self-compatibility of the hybrids, either by further backcrossing to self compatible S. eurylepis or continued inbreeding and the possible segregation for inbreeding.Other untested hypotheses are that S. chaseaewas derived from one or the other parent, or that S. chaseae might have given rise to the other two taxa (e.g., 1, EURY- 3 CHAS-- FOET; 2, FOET-- CHAS -* EURY; 3, EURY 4-CHAS-- FOET). Hypotheses 1 and 3, however, suggest that a self-compatible species gave rise to a self-incompatible one, the reverse of the usual situation in plantswhere inbreeders usually are derived from outbreeders (Grant 1981;Brauner & Gottlieb 1987). In addition, the apparently rearranged chromosomes of S. chaseae and S. eurylepis in relation tomost of the rest of Simsia (Table 4) suggest that these species are derived. Recent studies using more precise molecular mark ers have both supported (Warwick et al. 1979; Rieseberg et al. 1990) and rejected (Rieseberg et al. 1988; 1990) diploid hybri-dspeciation hypotheses. Simsia chaseae should be subjected to suchmolecular tests to provide more reliable data concern ing its origin. Simsia chaseae shows morphological similarity to S. dombeyana of South America (see Numerical Taxonomy), which suggests the close relationship of these two species, but the nature of their relationship is presently unknown. Conclusions. The foregoing hybridization data indicatemuch genetic and chro mosomal similarity for most of the species of Simsia. The data show that flowering F1 hybrids can be produced inmany wide combinations within the genus. Lowered crossability is correlated with my intuitive ideas of relationships, derived indepen dently of the crossing data. The artificialF1 hybrids aremorphologically intermediate between the parents in many features, and the taxonomic difficulty of some naturally-occurring taxa (e.g., S. amplexicaulis intergradingwith S. foetida var. foetida, and the intergradation of some of the varieties of S. foetida) possibly is caused by natural hybridization. The intergradation and intermediacyof some taxa, and the genetic and chromosomal similarity found throughout the genus suggests a recent origin of many of the species. The largely allopatric distributions of most of them suggest initialgeographic isolation and only partial divergence after separation.

MORPHOLOGY

Duration and Habit. Simsia comprises annuals and perennials. Some annuals (e.g., S. foetida, S. lagascaeformis) occasionally persist from a woody base, espe cially inwetter sites (see discussion under S. lagascaeformis).The perennials grade from herbaceous perennials to suffrutescent perennials (e.g., S. sanguinea, S. setosa) to shrubs (e.g., S. fruticulosa). All species usually have erect or ascending branches. Simsia amplexicaulis, S. lagascaeformis, and S. sanguinea occasionally are decumbent. Maximum plant height varies from about 1.5 m (e.g. S. calva, S. tenuis) to ca. 4 m, for both annuals (e.g. S. foetida var. grandiflora) and perennials (e.g., S. fruticulosa, S. steyermarkii). Roots. Simsia calva is the only species of Simsia with a fusiform-thickened, somewhat fleshy root. All other species have narrower taproots. Simsia am 1990 SIMSIA 17

plexicaulis and S. lagascaeformis often root at the lower nodes when their stems are decumbent. Leaves and Nodes. In all species, three to five main veins arise from the base of the blade at a point considered the junction of the petiole and blade. Some species (e.g., S. chaseae, S. dombeyana), have entirely unwinged petioles and nodes; oth ers (e.g., S. amplexicaulis, S. foetida var. megacephala) commonly have petioles partly to entirely winged to the base, often with amplexicaul bases. This character can vary on different plants in a population or on individualplants. Sometimes just the base of the petiole iswinged; these basal wings are referred to as stipules. Other species (e.g., S. setosa, S. villasenorii) commonly have nodal discs that are variously developed, from minute to 3 cm or more long and wide. The nodal disc is a structure connecting the bases of adjacent petioles across the nodes, or sometimes is reduced to a flap of tissue between the bases of adjacent petioles. Simsia annectens commonly has perfoliate leaves, but some individuals have the winged petioles tapering to the base and possess nodal discs. The extensive variability of all of these conditions occasionally can make identifications difficult, and in some instances itmay be necessary to examine more complete specimens or more than one plant from a population to use the keys effectively. Compositae generally are regarded as lacking stipules (Hutchinson 1973; Cronquist 1981). The term stipule commonly refers to an appendage, or pair of appendages, or sometimes a sheath, attached to the petiole or free and attached to the nodes (Fahn 1982; Howard 1979). Such structures occur within Simsia, else where in Helianthinae (e.g. Axiniphyllum Bentham, Turner 1978; Tithonia, La Duke 1982), in other subtribes of Heliantheae (e.g., Milleria L., Stuessy 1975, Sigesbeckia L., D'Arcy 1975), and inLiabeae (Nordenstam 1977;Robinson 1983), where they are referred to as "pseudo-stipulate petioles" or "stipule-likewings" or "nodal discs". These structures in Simsia and in these other genera may or may not be homologous with stipules of other dicotyledons. They are referred to as stipules or nodal discs here for simplicity of description. The blade outline of most species is ovate to deltate. The apices are acute to acuminate, and the bases cordate to truncate to attenuate. Simsia santarosensis has elliptic to ovate leaves, and S. sanguinea has some populations with linear leaves. Some species have leaves that are never lobed (e.g., S. santarosensis, S. vil lasenorii), others have leaves usually unlobed but rarely lobed (e.g., S. foetida, S. lagascaeformis), and still others have leaves usually lobed and occasionally unlobed (e.g., S. amplexicaulis, S. fruticulosa). Sometimes these latter two species have lobed and unlobed leaves present on the same individual. Leaf odor also is taxonomically useful. Simsia chaseae, S. foetida, and S. setosa are characterized by a range of spicy or lemony odors that are lacking in all other species. Vesture. Vesture is extremely variable inmany species, but sometimes can be taxonomically useful. There are six basic hair types in Simsia; unicellular, uniseriate multicellular (Figs. 6, 7), uniseriate with a complex multicellular base (Fig. 6), biseriate (Zwillingshaare, Hess 1938), uniseriate glandular (Fig. 8), and biseriate glandular (Fig. 9). The vesture may be scabrous, hirsute, hispid, strigose, pubes cent, puberulent, pilose, or sericeous (terminology followingHarrington and Dur rell 1957). The unicellular hairs taper to the apex except for one type that is blunt (Fig. 9). This blunt hair is universally present in all taxa on leaves, stems, and 18 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 30

FIG. -11 Pbecene ype ad tyl bancesinSimiaandViuira.6 dobean (Spooner2907),adaxialleaf lamina with glandular-puberulent and scabrous vesturewithalarger~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~Al

unseitemltcllla ai it mliclllr ae.7S haee Sooe 20),peieljstblo hea shwiggandla-puerlen an srigsevesure8 dobeyna(Sponr 207)susesi

uniseriate gland. 9~~~~~~~~~~~~..S. fo.d. ar.o.da(poer&Badebr.30).tpiaebseit.lad.n

smaller curved blunt-tipped unisriate hairs. 10. V cordifolia (pooner & Brandenburg.2314) shor stylebranches withdeltoid tips.dombeyana 11.S. (Spooner 2907) ofapexfihiform style branch~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~...... 1990 SIMSIA 19

peduncles. The biseriate hairs are confined to the surfaces of the achenes and are invariant in the genus. Glandular hairs (glands) are common in the genus. Uniseriate glands are subsessile andmost commonly found on the laminas.Biseriate glands are subsessile to stipitate. They are most common on the laminas and peduncles. Stipitate, biseriate glands are present in some taxa (e.g., S. foetida var. foetida, var. megacephala). These glands help to distinguish these varieties from S. foetida var. grandiflora, var. jamaicensis, and var. panamensis, which possesses subsessile, biseriate glands, but there is some intergradation. Glandular hairs of some type generally are present on the leaves of most species. Simsia amplexicaulis, S. calva, S. lagascaeformis, and S. santarosensis, however, generally lack leaf glands but sometimes have them. Uniseriate and subsessile biseriate glands often have the superficial appearance of sessile resin droplets under binocular optics and often are difficult to distinguish from each other. Because of the mixed distribution and gradation of these types, designations are not cited in the descriptions except for the varieties of S. foetida, where they are diagnostic. .All species have corymbiform or paniculate (tech nically capitulescences) arising from the distal nodes of terminal and lateral branches; S. calva and occasionally S. foetida are monocephalous. In some species, the peduncle length and associated aggregated nature of the inflorescence are taxonomically useful, as in S. steyermarkii,with its tightly aggregated inflorescence, or S. fruticulosa, in whicih the heads are tightly to moderately aggregated. Many other species (e.g., S. annectens, S. eurylepis, S. villasenorii) are variable in this feature. The inflorescences usually have moderately divaricately diverging peduncles, but some individuals of S. annectens commonly have peduncles that are widely divaricately diverged. This feature is highly variable within this species and of limited taxonomic utility. Heads. Simsia eurylepis is discoid; all other species are radiate. Reports of discoid heads (e.g., S. chaseae,Millspaugh 1904, S. dombeyana [as S. hirsutaBlake 1913]Burkart 1974; Cabrera 1978, S. lagascaeformis [as S. exaristata,Blake 1913]) are possibly errors. Occasionally, short ligules are included or little exserted beyond the inner phyllaries of these three latter species. Head size and the associated numbers of phyllaries and florets are taxonomi cally useful for separating the varieties of S. foetida; var. grandifiora and var. panamensis are distinguished from other varieties by large numbers of disc florets (66-172 and 40-151, respectively). Also, S. tenuis is partly distinguished from related species by its small heads containing 12-14 disc florets. All head diameters in the descriptions include the extended rays. Head shape during flowering and early fruiting grades from campanulate to ovoid-campanulate to urceolate. For example, the large campanulate heads of S. foetida var. grandiflora and var. panamensis grade into the more ovoid-campanulate heads typical of var. foetida. The three self-compatible species (S. chaseae, S. dom beyana, and S. eurylepis) tend to have urceolate heads. The difference between ovoid-campanulate and urceolate is often difficult to distinguish on flattened herbar ium specimens. Involucre. The involucre is composed of 11-66 imbricate phyllaries in two to four series. The number of series varies and often is related to size of the head; larger heads have four series. As a result of this variation, this character is not taxonomically useful in the genus. There are two interpretations concerning the 20 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 30

identity of that subtend the ray florets. Stuessy (1977) regards them as part of the involucre. Funk (1982) regards them as pales inMontanoa Cerv. because of their similar development with other pales during fruiting. In Simsia the bracts are flat and herbaceous, and are similar to the other involucralbracts and therefore are here regarded as part of the involucre. The ultimate orientation of the phyllaries varies in different taxa from appressed (e.g., S. eurylepis) to the tips reflexed (e.g., S. annectens var. annectens), but there ismuch variation elsewhere. For example, S. calva has some individualswith the phyllary tips greatly reflexed (Spooner & Schil ling 2385) and others with the phyllary tipsmore appressed (Spooner & Schilling 2391). Four aspects of the involucre have much more taxonomic utility: phyllary grada tion (relative lengths of the different series), margin, color, and apex. Some species (e.g., S. lagascaeformis, S. villasenorii) are characterized bymiddle and outer series of phyllaries progressively shorter than the inner series (gradate). In other species (e.g., S. dombeyana, S. foetida), all series are subequal to each other in length. Simsia amplexicaulis and S. sanguinea have phyllaries of intermediate gradation, but the former tends toward subequal phyllaries and the latter towardmore gradate ones. The phyllaries of most species are almost always entire, but those of S. foetida (and rarely S. calva and S. annectens var. annectens) have a single sinuate constric tion near the apex. Phyllary color ranges from yellow to green and black to purple, with mottling (as green speckled with black) or gradations of color present on the same phyllary. Purple is found only in certain species. The apex of the phyllaries is acute to acuminate, but in S. foetida, S. amplexicaulis, and S. annectens var. annectens is commonly caudate. Pales. Most species possess conduplicate, persistent to tardily deciduous pales of medium texture. The pales are wide at the base and taper to an acute or acuminate apex (e.g., S. villasenorii). Simsia lagascaeformis has some populations in which the pales are subtruncate at the apex, and S. foetida tends to have pales with linear-acuminate apices. The vesture is generally antrorsely pubescent, but in S. foetida tends to be strigose. Color varies from totally stramineous to variously flecked with black or purple, especially apically, and some populations of some taxa have pales that are entirely purple. The pales often are narrower in the center of the head and sometimes are reduced to linear bracts. All of the above features are highly variable. One pale character found to have more stability is the slightly longer length of pales of S. foetida var. megacephala (often slightly exserted beyond the disc florets), which helps distinguish this variety from others of S. foetida. Ray corollas. Ligule color is taxonomically useful, and ranges from pale yellow or light lemon-yellow (e.g., S. foetida var. foetida, S. villasenorii) to light orange yellow (e.g., S. fruticulosa, S. molinae), to darker orange-yellow (e.g., S. am plexicaulis, S. lagascaeformis), or may be pink to red to deep purple or rarely white (S. sanguinea). Although there are intermediate yellow hues, the pale or lemon yellow taxa are relatively uniform (except in S. foetida var. grandiflora and S. foetida var. panamensis in which most populations are orange-yellow, but some grade to lemon-yellow). The intermediate yellows of these two varieties appear closer to orange-yellow. Although ligule colors are easily discerned in the field, they are harder to distinguish on herbarium specimens. Discflorets. Taxonomically useful characters are found in the length of the tube of the disc corolla, the color of the anther thecae, the color of the terminal anther 1990 SIMSIA 21

appendage, and the length and shape of the style branches. The relatively longer tubes of S. foetida var. jamaicensis help to distinguish this variety from the others within S. foetida, but these differences in length have limited utility elsewhere. Anthers. Anther thecae vary from yellow to brown to black; Simsia ghies breghtii always has black thecae. Many other species, polymorphic for this char acter, have anthers that are predominantly black or brown but occasionally yellow (e.g., S. fruticulosa, S. setosa). Simsia amplexicaulis, S. lagascaeformis, and some times S. sanguinea have thecae that are yellow basally but purple (often drying bronze) on the distal V3-?2. Simsia sanguinea almost always has black or black rimmed terminal anther appendages with the black color sometimes extending to the apex of the thecae. In all other species, terminal anther appendages are stramineous (or green turning stramineous). The long (2-4 mm) and thin style branches with an attenuate apex provide a useful generic character that helps to distinguishmost species of Viguiera (Fig. 10) from Simsia (Fig. 11), but the styles of Simsia and some members of Viguiera ser. Grammatoglossae are similar (see Taxo nomic History and Generic Relationships). Achenes. The achenes of all species are very similar and provide major charac ters that define the genus. All species have unwinged achenes that exhibit extreme lateral flattening. Some populations of S. ghiesbreghtii have unwinged but laterally thickened, biconvex achenes similar to those of some species of Viguiera (e.g., V. ovata Blake). Achenes of Simsia are very similar in outline, varying from broadly ovate to suborbicular, but much intraspecific variation of these shapes precludes their use taxonomically. The apex of the achene varies from subtruncate and slightly concave to notched apically with well-developed apical shoulders, but this trait also varies within taxa. In size, the achenes vary from relatively small in S. ghiesbreghtii (2.6-3.5 mm long, 1.2-1.9 mm in diameter) to relatively large in S. foetida var. jamaicensis, var. megacephala, and S. dombeyana (4.2-7.5 mm long, 2.5-4.1 mm in diameter). Much similarity and overlap of these sizes precludes taxonomic utility except in the cases of S. foetida var. panamensis, S. ghiesbreghtii, and S. steyermarkii,whose relatively small achenes help to distinguish them from related taxa. Pappus. Most achenes are awned and pubescent. These awns are easily de tached and often tardilydeciduous. Many species are known to have some individu als with calvous (bald, hairless, glabrous; Stearn 1983) achenes. Rarely, epappose and pubescent or awned and glabrous achenes are encountered. and S. F. Blake placed much taxonomic weight on these achenial characters. Gray (1846) originally described Barrattia partly based on its calvous achenes, but later (Gray 1850) realized the tenuous nature of this trait and combined Barrattia and Simsia. Presence/absence of awns and/or vesture of the achenes also have been used to distinguish species (S. calva vs. S. subaristata, Gray 1849; S. ghiesbreghtii vs. S. sericea, Gray 1883; Blake 1913; Robinson & Brettell 1972; S. exaristata vs. S. lagascaeformis, Gray 1853; Blake 1913), and varieties (within S. amplexicaulis, Blake 1913, 1917; S. calva, Blake 1913; S. exaristata, Blake 1917). Variability of these characters previously has been known to occur in S. setosa and S. sanguinea (Blake 1917). My investigations document intrapopulational variability of these characters in the following populations: S. amplexicaulis, Spooner & Villasenior 2557; S. annectens var. annectens, Spooner 2845; S. calva, Spooner & Schilling 2385; S. chaseae, Spooner 2803; S. dombeyana, Jorgensen 1286; S. eurylepis, 22 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 30

Spooner & Brandenburg 2209; S. lagascaeformis, Spooner & Burgos 2602; S. san guinea, Spooner & Burgos 2604, Spooner & Villasefior 2505, 2567; S. setosa, Spooner & Villasefior 2528; and S. steyermarkii, Spooner & Dorado 2748. The specimen of S. dombeyana mentioned above has awns minute to fully formed on different achenes of the same head. In addition, I obtained both awned and non awned progeny from one individual of S. eurylepis (Spooner & Brandenburg 2209) raised from an awned achene. Vesture and presence/absence of awns on the disc achenes is too variable to be taxonomically useful at any level within the genus. Squamellae (scales, usually short and lacerate, attached to the apex of the achene between the awns) rarely are borne by achenes of some species of Simsia. Formerly, their absence within the genus had been used inpart to distinguish Simsia from Viguiera, in which they are usually present. The achenes of S. ghiesbreghtii often bear 4-12 squamellae, and those of S. villasenorii often four. All other species commonly lack them, but four squamellae have been observed on some achenes of the following: S. dombeyana, Haught 249; S. foetida var. foetida, Hinton 5538; S. sanguinea, Spooner & Villasefior 2570. They are similar to squamellae of many species of Viguiera and perhaps are homologous structures.

CONCEPTS OF SPECIES AND VARIETIES

Simsia presents many problems in the circumscription of species because of intermediates and extensive variability of characters. The species concept adopted here is based entirely on morphology. Species are here considered "the smallest groups that are consistently and persistently distinct, and distinguishable by ordi narymeans" (Cronquist 1978), although occasional intermediates sometimes pro vide taxonomic difficulties. Varieties are designated instead of subspecies, because I wish to use a lower taxonomic rank for very similar taxa. Subspecies and varieties frequently are used interchangeably when only one intraspecific category is employed (Kapadia 1964). I regard varieties as distinct but often intergrading sets of populations with a geo graphic component. Numerous other variations of many features in Simsia could formally be recognized as formae, but I agree with Burtt (1970) that such formal recognition only serves to impose unnecessary complexity on a taxonomic system. These variants are best discussed but left unnamed.

TAXONOMY

Morphological similarity and intergradation among some taxa of Simsia (e.g., the varieties of S. annectens and the varieties of S. foetida) sometimesmake specific delimitations complex and attempts at identifications difficult. Frequently, taxa can be recognized only by a combination of character states, including some overlap ping ones (e.g., S. chaseae vs. S. dombeyana). Polymorphic character states present the greatest problem in key construction (e.g., differences in ligule colors of S. sanguinea or another color in many species). Nodal morphology can be especially difficult. Many species are distinguished by nodal discs, but some nodes or occa sional individuals in a population lack nodal discs, and some species usually lacking nodal discs possess them. Attempts to include all possible exceptions in the keys 1990 SIMSIA 23 would make them unworkably cumbersome. Also, some features such as plant height, leaf texture, or ligule color are not always easily obtained from herbarium specimens, but these characters are evident and diagnostic on living specimens.

Simsia Persoon, Syn. pl. 2: 478. 1807, non Simsia R. Brown, 1810. Encelia sect. Simsia (Persoon) A. Gray, Proc. Amer. Acad. Arts 8: 656. 1873. LECTOTYPE,designated by Robinson and Brettell, 1972: Simsia amplexi caulis (Cavanilles) Persoon. Armania Bertero ex de Candolle, Prodr. 5: 576. 1836.-TYPE: Armania fruticulosa (Sprengel) Bertero ex de Candolle. [=Simsia fruticulosa (Sprengel) S. F. Blake]. Barrattia A. Gray & Engelmann ex A. Gray, Proc. Amer. Acad. Arts 1: 48. Dec 1846 or Jan 1847 [fide Stafleu and Cowan, 1976].-TYPE: Barrattia calva A. Gray & Engelmann ex A. Gray [=Simsia calva (A. Gray & Engelmann ex A. Gray) A. Gray].

Erect or ascending, taprooted, annual or suffruticose perennial herbs or shrubs (decumbent and rooting at the nodes in some populations of S. amplexicaulis and S. lagascaeformis,with roots fusiform-thickened in S. calva). Stems sparingly to freely branched, terete to slightly ridged, yellow to green to purple, glabrous to puberulent to scabrous or hispid, glandular or eglandular. Leaves opposite and decussate below (rarely whorled), alternate above, those on successive orders of branching or on the upper parts of the stem often reduced in size and becoming -like in the inflorescence; petioles without wings to partially or completely winged (perfoliate on some nodes of S. annectens), often with expanded bases of adjacent leaves fused to form nodal "discs" that are fused to the stem and bases of adjacent petioles or reduced to interpetiolar structures, sometimes deciduous; petioles present or absent, vesture usually as on the blades (often long-pilose at base in S. eurylepis and S. lagascaeformis);blades 3-nerved from the base, cordate to linear, unlobed to deeply 3-5-lobed (pinnatifid in some populations of S. sanguinea), thin and herbaceous to thick and stiff, glabrous to scabrous to sericeous, glandular to eglandular, margins crenulate to coarsely dentate or rarely entire. Inflorescence of solitary heads or a tightly to loosely branched corymb or panicle, the peduncles sometimes divaricately branched; peduncles usually glandular-puberulent, often scabrous to strigose or hispid. Heads heterogamous, radiate (eradiate in S. eurylepis). Involucre campanu late to ovoid-campanulate to urceolate, base truncate or indented at the attachment to the peduncle. Receptacle low-convex. Phyllaries 11-66, tightly appressed to broadly reflexed, acute to acuminate to caudate at the tips, green to yellow, purple, or blackish, scabrous to hispid to sericeous, glandular or eglandular, ciliate at the margins, 2-4-seriate, imbricate, unequal with the middle and outer series progres sively shorter than the inner or all phyllaries subequal, the outer series ovate to linear, entire (with a single sinuate constriction at the margin in S. foetida, and rarely in S. calva or S. annectens var. annectens); inner phyllaries lance-ovate to linear, entire to sinuate. Pales radially conduplicate and enclosing the achenes, of medium texture, stramineous to purple or black, often mottled in color only at the apex, sometimes colored throughout, glabrous or pubescent along the margins and midrib, especially apically. Ray florets (0-) 5-45, in a single series, sterile; ligules light lemon-yellow to orange-yellow (various hues of pink to purple or rarely white in S. 24 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 30 sanguinea), entire to 2-3-lobed at apex, glandular-puberulent; ovaries linear, slightly tapering from the base to the apex, trigonous, with or without awns. Disc florets 12-172, perfect; corolla concolorous with the rays (orange-yellow in eradiate S. eurylepis, white inmost populations of S. sanguinea), usually turning purple apically; tube glandular-puberulent, abruptly expanded above to a subcylindric throat; throat ventricose and sparingly glandular-puberulent at the base, slightly flared and glabrous apically; lobes 5, triangular, sparingly to densely pubescent, sometimes glandular on the outer surface, papillose on the inner surface; stamens 5, anthers yellow to brown to black but sometimes darkened only apically (usually yellow proximally and bronze or purple distally in S. amplexicaulis and S. lagascae formis, sometimes so in S. sanguinea), glabrous ormore commonly antrorsely pubes cent along the connectives, terminal appendages stramineous (black or black rimmed in S. sanguinea); style branches slender, 2-4 mm long, with or without an attenuate apex; achenes light brown to black or mottled, laterally flattened (thick ened and biconvex in some populations of S. ghiesbreghtii), transverselyobovate to elliptic in outline with minute to conspicuous apical shoulders extending beyond the attachment of the corolla, glabrous or hirsute with double hairs; pappus absent or of two ascending awns often flared basally, rarelywith 4-12 shorter intermediate squa mellae. Chromosome number: n = 17.

KEY TO THE SPECIESAND VARIETIES OF SIMSIA

1. Root fusiform-thickened; heads borne singly on peduncles arising from each of many nodes; rays yellow, often brown or purple-lined abaxially. 12. S. calva. 1. Root fibrous or a taproot, but not fusiform-thickened; heads aggregated on peduncles branched past the nodes; rays yellow, pink to deep purple, or white, always the same color ab and adaxially. 2. Rays pink to deep purple, rarely light lemon-yellow or white; terminal anther appendages usually black or stramineous and black on margin; petioles usually winged, nodes without discs. 11. S. sanguinea. 2. Rays light lemon-yellow to orange-yellow; terminalanther appendages stramineous through out; petioles winged or unwinged, nodes with or without discs. 3. Phyllaries subequal in length, the outer series usually ? or more the length of the inner. 4. Leaves perfoliate or with conspicuous nodal discs, the discs usually attached to the bases of the petioles. 5. Leaves sericeous abaxially; achenes 2.6-3.5 mm long. 1. S. ghiesbreghtii. 5. Leaves scabrous and hirsute abaxially; achenes 3.5-7 mm long. 7a. S. annectens var. annectens. 4. Leaves not perfoliate; nodal discs, if present, generally confined to the nodes and not attached to the base of the petioles. 6. Leaves mostly eglandular; anther thecae usually yellow proximally, commonly purple or bronze distally. 17. S. amplexicaulis. 6. Leaves densely glandular; anther thecae yellow or black, never purple distally. 7. Involucres urceolate. 8. Outer phyllaries 1.6-3.1 mm wide; inner phyllaries 1.2-2.9 mm wide; South America. 16. S. dombeyana. 8. Outer phyllaries 1.1-2 mm wide; inner phyllaries 1.2-2.2 mm wide; Gulf coastal lowlands of Mexico. 15. S. chaseae. 7. Involucres campanulate to ovoid-campanulate. 9; Leaves pilose to subsericeous on both sides with intermixed glandular puberulent vesture; achenes 3.1-5.1 mm long; Panama and South America. 18e. S. foetida var. panamensis. 1990 SIMSIA 25

9. Leaves glandular-puberulent and setose to hispid on both sides with inter mixed glandular-puberulent vesture; achenes 3.5-7.5 mm long;Mexico to Costa Rica or Jamaica. 10. Ligules orange-yellow, sometimes lemon-yellow; heads 25-45 mm in diameter, phyllaries 21-66; disc florets 66-172; Guatemala to Costa Rica. 18d. S. foetida var. grandiflora. 10. Ligules lemon-yellow; heads 20-35 mm in diameter, phyllaries 11-29; disc florets 20-76; Mexico to or Jamaica. 11. Petioles of the distal part of the plant usually winged throughout, the wings often 1 cm wide or more, the proximal nodes with stipules to 3 cm long; pales 9.2-14 mm long, frequently equalling to slightly longer than the disc florets; eastern Oaxaca. 18c. S. foetida var. megacephala. 11. Petioles of the distal part of the plant unwinged or with wings gener ally less than 5 mm wide; proximal nodes with stipules generally less than 1 cm long; pales usually shorter than disc florets. 12. Leaves with stipitate glands; tube of disc floret 0.6-1.8 mm long;Mexico, Guatemala, El Salvador. 18a. S. foetida var. foetida. 12. Leaves with subsessile glands; tube of disc floret 1.7-1.9 mm long; Jamaica. 18b. S. foetida var. jamaicensis. 3. Phyllaries unequal in length, themiddle and outer series progressively shorter, the outer series 2 or less the length of the inner. 13. Heads eradiate. 14. S. eurylepis. 13. Heads radiate. 14. Anthers yellow, usually purple or bronze distally; nodes sometimes long-pilose; phyllaries usually purple. 13. S. lagascaeformis. 14. Anthers yellow to brown to black throughout; nodes glabrous to scabrous or puberulent or pubescent but not long-pilose; phyllaries purple to green or green black. 15. Nodal discs absent; involucres urceolate; Gulf coastal lowlands of Mexico. 15. S. chaseae. 15. Nodal discs usually present (absent in S. santarosensis); involucres campanu late to ovoid-campanulate; Mexico, , and South America. 16. Leaves subsessile or with petioles to 1 cm long, without nodal discs, blades stiff and unlobed. 2. S. santarosensis. 16. Leaves distinctly petiolate, petioles 0.5-11 cm long, nodal discs present at most nodes, blades stiff to herbaceous, lobed or unlobed. 17. Leaves stiff and thick, unlobed. 9. S. villasenorii. 17. Leaves herbaceous, lobed to unlobed. 18. Leaves commonly perfoliate, but some petioles winged only partly to the base. 7b. S. annectens var. grayi. 18. Leaves not perfoliate, with petioles generally unwinged, except at the very base where attached to the nodal disc. 19. Leaves sericeous abaxially (sometimes pilose in S. fruticulosa). 20. Achenes laterally flattened to biconvex, 2.6-3.5 mm long, 1.2-1.9 mm wide, often with awns and squamellae, sometimes epappose; Chiapas, Mexico to Guatemala. 1. S. ghiesbreghtii. 20. Achenes laterally flattened, 4-6.4 mm long, 1.7-3.0 mm wide, awned but without squamellae; South America. 3. S. fruticulosa. 19. Leaves puberulent, pilose, hispid, or hirsute abaxially. 21. Disc florets 12-14; annuals or suffrutescent perennials. 8. S. tenuis. 21. Disc florets 15-85; suffrutescent perennials or shrubs. 22. Phyllaries usually purple, sometimes green; ligules 5-7 mm long; anthers yellow. 5. S. holwayi. 26 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 30

22. Phyllaries green to green-black; ligules 7.5-19.5 mm long; anthers yellow to brown or black. 23. Heads tightly aggregated, or subsessile, or on peduncles to 2 cm long; achenes 3.2-4.2 mm long, 1.8-2.5 mm wide. 6. S. steyermarkii. 23. Heads in open inflorescences, peduncles to 7 cm long; achenes 4.2-6.5 mm long, 2.5-4 mm wide. 24. Stems glandular-puberulent, strongly his pid and hirsute; ray florets light lemon yellow; northwesternMexico. 10. S. setosa. 24. Stems puberulent to pubescent, weakly hispid; ray florets lightorange-yellow; Cen tralAmerica. 4. S. molinae.

1. Simsia ghiesbreghtii (A. Gray) S. F. Blake, Proc. Amer. Acad. Arts 49: 392. 1913. Encelia ghiesbreghtii A. Gray, Proc. Amer. Acad. Arts 8: 658. 1873.-TYPE: MEXICO.Chiapas: mountain forest near Chiapas, 1864-1870, Ghiesbreght 568 (holotype: GH!). Verbesina argenteaBertoloni, Novi Comment. Acad. Sci. Inst. Bononiensis 4: 435. 1840, non Verbesina argenteaGaudichaud, 1829.-TYPE: GUATEMALA. Sacatepequez: "Vulcano d'acgua" [=Volcain de Agua], 1837, Vallas[quez] [=Valasquez, fideBlake, 1926] s.n. (holotype: BOLO, photo: OS!). Simsia sericea (Hemsley) S. F. Blake, Proc. Amer. Acad. Arts 49: 393. 1913. Encelia sericeaHemsley, Biol. cent.-amer., Bot. 2: 185. 1881.-TYPE: GUA TEMALA.Montagua Valley, 1861, Salvin & Godman 133 (holotype: K!, fragment:US!, photos: OS! US!).

Suffrutescent, taprooted perennials to shrubs. Stems 1-2 m tall, 0.3-2 cm in diameter, erect, green to purple to brown, puberulent to pilose. Leaves with peti oles 1-3 cm long, 1-2 mm in diameter, at base with stipules of opposite leaves fused to form nodal discs 0-1.5 cm long and wide; blades 4-15 cm long, 1.5-8 cm wide, ovate to deltate, herbaceous, unlobed, base cordate to truncate to cuneate, apex acute to acuminate, eglandular, puberulent or hispid to pubescent adaxially, sericeous abaxially, margins subentire to crenate to dentate. Inflorescence corymbi form, loosely branched; peduncles 1-6 cm long, 1 mm in diameter, puberulent, sometimes intermixed short-pubescent and glandular. Heads radiate, 10-15 mm long, 20-40 mm in diameter. Involucre campanulate, 8-13 mm long, 7-11 mm in diameter; phyllaries 15-20, 2-3-seriate, generally unequal, sometimes subequal, usually appressed except slightly reflexed apically or less commonly greatly re flexed, dark green to green-black, puberulent to pilose, rarely glandular, margins entire; outer phyllaries 3.6-7.7 mm long, 1-2 mm wide, lance-ovate; inner phyllaries 6.7-12 mm long, 1.5-2.5 mm wide, lanceolate. Pales 5.5-8.8 mm long, 1.5-2.3 mm wide, stramineous, green-black apically, acute to aristate, strigose to antrorsely short-pubescent and sometimes glandular apically. Ray florets 7-11; ligules 8-15 mm long, 3-6.6 mm wide, light orange-yellow; tube 1-1.7 mm long, 0.3-0.6 mm in diameter; ovary 3-4.5 mm long, 0.4-1 mm wide. Disc florets 31-50; corolla 4.9-6.7 mm long, 0.9-1.5 mm in diameter; tube 0.7-1.6 mm long, 0.3-0.8 mm in diameter; lobes 0.7-1.4 mm long, 0.5-0.8 mm wide; anthers 3-4.2 mm long, black, terminal appendages stramineous; achenes 2.6-3.5 mm long, 1.2-1.9 mm 1990 SIMSIA 27

900

200~~~~ .8 . 20

S. ghie,br i I I..x ...... i...... 2

I...., XI I

x~~~~IS.hebehi

A S. holwayi * S. molinae * S. santarosensis * S. steyermarkii

900

FIG. 12. Distribution of Simsia ghiesbreghtii, S. holwayi, S. molinae, S. santarosensis, and S. steyermarkii.

wide, laterally flattened to laterally thickened and biconvex, awns absent or 1-2.7 mm long, squamellae absent or 4-12, 0.4-0.7 mm long. Chromosome number: n = 17. Phenology. Flowering fromOctober throughApril. Distribution (Fig. 12). In the northern highlands of Chiapas, Mexico, and along the volcanic belt in Guatemala; in openings, on steep wet mountain slopes and occasionally in dry roadside thickets; 1600-2500 m.

ADDITIONAL SPECIMENS EXAMINED. [Note: All collections reported only from "Volcan de Agua," including the type of Verbesina argentea, are listed under the department of Sacatepequez, although the slopes of this volcano also are situated in the departments of Escuintla and Guatemala]. Mexico. CHIAPAS: near Navenchauk, Breedlove 9507 (DS, MICH), 48485 (CAS, OS), Spooner 2781 (COL, MEXU, OS, TENN, WIS). Guatemala. BAJA VERAPAZ: near Pantin on rd to Salama, Spooner & Linder 2753 (MEXU, OS, TEX), Williams et al. 42168 (BM, DUKE, F), Williams et al. 43367 (F, MICH).-EL PROGRESO: between Calera and middle slopes of quebradas of Volcan Siglo, Steyermark 43018 (F, GH).-EL QUICHE: 10 mi NE of Sacapulas, Williams et al. 41210 (F, NY).-GUATEMALA: San 28 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 30

Rafael, collector unknown 63 (GH).-QUEZALTENANGO:E slope of Volcdn SantaMaria de Jesus, just S of Quezaltenango, Spooner & Dorado 2675 (MEXU, OS, OSH, RSA, US, WIS), Stuessy & Gardner 4345 (ASU, ENCB, OS).-SAN MARCOS:between San Marcos & Castalia, Spooner & Dorado 2674 (MEXU, OS, P, RSA, US, WIS), Williams et al. 26142 (F, GH, NY, US).-SACATEP9QUEZ:Antigua, Kellerman 4982 (OS, US); Volcin de Agua, Feb 1905, Kellerman s.n. (US), Maxon & Hay 3751 (US), Molina 21030 (F, NY), Molina & Molina 24895 (BM, F, S), 24913 (C, DUKE, F, NY, US), Standley 59379 (CAS, F, MICH, U), 65054 (F); Finca El Hato, NE of Antigua, Standley 61158 (F, US), 61199 (F, U); 10 km from San Mateo del Milpar, Molina 15512 (F, GH, US).-SOLOLA: 8 mi E of Panajachel, Spooner & Dorado 2683 (LE, MEXU, OS); 5 km E of Panajachel above Lake Atitlan, Williams et al. 25231 (F).

Simsia ghiesbreghtii is distinguished by its sericeous leaves and relatively small achenes, which are the smallest in the genus. It is superficially similar to S. fruticulosa, which also has sericeous leaves. These species are readily distinguished by the achenes; those of S. fruticulosa are 4-6.6 mm long and never laterally thickened, whereas those of S. ghiesbreghtii are 2.8-3.5 mm long and sometimes laterally thickened. Simsia fruticulosa is often a much largerplant, growing to 4 m tall,whereas S. ghiesbreghtii grows to only 2 m tall. Simsia ghiesbreghtii exhibits great variation in achene morphology, but this is not taxonomically useful within the species. As inmany other species of Simsia, there is intraspecific variation for presence or absence of indument and pappus, a feature that is insignificant taxonomically throughout the genus (see Morphology and Taxonomic Criteria). Previously, S. ghiesbreghtii has been distinguished from S. sericea by the calvous achenes of the former. When a pappus is present, it is of two awns or of two awns and 4-12 intermediate squamellae. A feature of greater taxonomic significance in S. ghiesbreghtii is achene shape. Some populations (e.g., Kellerman 4982,Molina & Molina 24913) have achenes that are laterally flattened, typical of Simsia. Others (e.g., Standley 59379, Steyermark 43018, Williams et al. 43367) have laterally thickened achenes with no shoulder, similar to some species of Viguiera (e.g., V. ovata). Other populations exhibit intermediate morphologies. The presence of typically flattened Simsia-like achenes in some populations of S. ghiesbreghtii, plus the presence of nodal discs, rare in Viguiera and common in Simsia, influencemy placement of this species in Simsia rather thanViguiera. This species is possibly transitional, linking these two genera. Also variable in S. ghiesbreghtii are the relative lengths and pubescence of the phyllaries, from almost subequal and subsericeous (e.g., Breedlove 48485, Kel lerman s.n. [Feb 1905]) to unequal and puberulent to pubescent (e.g., Maxon & Hay 3751, Standley 59379). Because these differences exhibit no geographic compo nent, and because most collections are intermediate, this variation is treated as taxonomically insignificant. The exact type localities for Simsia ghiesbreghtii and Simsia sericea are un known. Hemsley (1887), in his brief account of the travels of the collectors of these types, did not resolve these questions. The Monagas Valley, the type locality of S. sericea, traverses the departments of Baja Verapaz, El Progreso, Zacapa, and Izabal. Hemsley (1887) commented that most of the collections of Salvin and God man came from Volcain de Fuego, a possible locality for this species, but this volcano is in the departments of Escuintla, Sacatepequez, and Solola. Likely locali ties for the type of S. sericea are in the departments of El Progreso or Zacapa in the Sierra de las Minas that borders the Monagas Valley to the north. 1990 SIMSIA 29

2. Simsia santarosensis D. M. Spooner, sp. nov.-TYPE: COSTARICA. Guanacaste: along unpaved rd, 0.5 mi N of El Gallo, N of Liberia, scrubby tropical deciduous forest, in whitish, sandy, volcanic soil, 29 Dec 1985, Spooner 2900 (holotype: OS!; isotypes: CR! F!MEXU! MICH! NY! OSH! TENN! UC! US! WIS!).

Plantae suffruticosae, 0.3-1.5 m altae, foliis sessilibus vel petiolatis petiolis usque ad 1 cm longis, estipulatis, laminis ellipticis vel ovatis, rigidis, non lobatis, 1 5 cm longis, 1-2 cm latis, basi cuneatis, apice acutis vel acuminatis, involucris gradatis. Suffrutescent, taprooted perennials. Stems 0.3-1.5 m tall, 0.3-1 cm in diame ter, erect, brown to purple, weakly strigose and canescent, becoming glabrous with age. Leaves subsessile or with petioles to 1 cm long, 1 mm in diameter, often cuneate, estipulate; blades 1-5 cm long, 1-2 cm wide, ovoid, stiff, unlobed, base cuneate, apex acute to acuminate, scabrous, mostly eglandular, but occasionally with very scattered glands, margins subentire-crenate. Inflorescence corymbiform, loosely branched; peduncles 1-14 cm long, 1mm in diameter, puberulent, some times scabrous and glandular. Heads radiate, 13-15 mm long, 20-25 mm in diame ter. Involucre campanulate, 11-13 mm long, 5-10 mm in diameter; phyllaries 19 29, 3-seriate, unequal, appressed or slightly reflexed apically, acute to acuminate at the tips, green to green-black, turning brown, canescent, sometimes scabrous, margins entire; outer phyllaries 3-6 mm long, 1.5-2.5 mm wide, ovate to lance ovate; inner phyllaries 8.5-11.5 mm long, 1.4-2.5 mm wide, lanceolate, linear, or oblanceolate. Pales 9-10.8 mm long, 1.6-2.2 mm wide, stramineous, acute to acuminate, antrorsely pubescent apically. Ray florets 8-13; ligules 10.5-11.5 mm long, 3.2-3.8 mm wide, light orange-yellow; tube 1-1.4 mm long, 0.4-0.6 mm in diameter; ovary 5.6-7.5 mm long, 0.4-0.7 mm wide. Disc florets 17-25; corolla 6 6.6 mm long, 1-1.5 mm in diameter; tube 0.7-1 mm long, 0.5-0.8 mm in diameter; lobes 0.9-1.3 mm long, 0.5-0.8 mm wide; anthers 3.3-4 mm long, yellow to black, terminal appendages stramineous; achenes 4.2-6.7 mm long, 2-2.7 mm wide, later ally flattened, awns 3.8-4.6 mm long, squamellae absent. Chromosome number: n - 17. Fig. 13. Phenology. Flowering from June through February. Distribution (Fig. 12). In the Pacific lowlands of Guanacaste, Costa Rica; in sandy volcanically derived soils; 0-400 m.

ADDITIONAL SPECIMENS EXAMINED. Costa Rica. Guanacaste: near Cainas, 11 mi S of Liberia, Daubenmire 493 (F); Comelco, Heithaus 168, 397 (MO); above Playa Naranjo, Santa Rosa National Park, Liesner et al. 2661 (MO); 4 km N of Liberia, Opler 574 (F).

This very distinctive species is easily recognized by its small, firm, elliptic to ovate, estipulate leaves. Only S. villasenorii has leaves as firm, but it has nodal discs, larger leaveswith longer petioles, and light lemon-yellow rather than orange yellow ligules.

3. Simsia fruticulosa (Sprengel) S. F. Blake, J. Bot. 53: 200. 1915. Hopkirkia fruticulosa Sprengel, Syst. veg. 3: 444. 1826.Armaniafruticulosa (Sprengel) Bertero ex de Candolle, Prod. 5: 576. 1836. Salmea fruticulosa (Sprengel) 30 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 30

sA ~~~~~~~~~~~c

FIG. 13. Simsia santarosensis (Spooner 2900). A. Stem with mature infloresence. B. Head. C. Pale. D. Disc floret with nearly mature achene. E. Ray floret.

Steudel, Nomencl. bot. 2: 775. 1841. Encelia fruticulosa (Sprengel) Hiero nymus, Bot. Jahrb. Syst. 19: 54. 1894.-TYPE: COLOMBIA.Magdalena: Santa Marta, [1820-1821 fide Urban, 1902], Bertero s.n. (holotype: P!, photos: GH! OS!; isotypes: B, destroyed, fragment:GH!, photos: GH! NY!, G-DC, microfiche: IDC. 800. 946. II. 4!). Simsia pastoensis Triana, Ann. Sci. Nat. Bot. ser. 4,9: 40. 1858.-TYPE: COLOM BIA. [Nariflo, fide Dugand, 1944; Pasto in protologue]: near Ortega, 1200 m, [1500m on label], 1851-1857, Triana 1362 (lectotype, here designated: P!, fragment: US!, photos: GH! OS! US!). Simsia pubescens Triana, Ann. Sci. Nat. Bot. ser. 4,9: 40. 1858.-TYPE: COLOM BIA. Cundinamarca: between Tena and El Colegio, 1400 m, 1851-1857, Triana 1361 (holotype: P!, fragments: F! US!, photos: GH! OS! US!; 1990 SIMSIA 31

isotypes: B, [destroyed, photos: GH! TEX!], BM! [photos:GH! US!, trac ing and fragment:GH!]). Encelia sodiroi Hieronymus, Bot. Jahrb. Syst. 29: 43. 1901. Simsia sodiroi (Hieronymus) S. F. Blake, Proc. Amer. Acad. Arts 49: 383. 1913.-TYPE: . Pichincha: dry rocky areas of the Rio Guallabamba, 18 Aug 1885, Sodiro 34/2 (holotype: B, destroyed, fragment: US!; isotype: Q!). Simsia grisea S. F. Blake, Contr. Gray Herb. 53: 53. 1918.-TYPE: VENEZUELA. Sucre: Guanta, 1 Jul 1917, Curran & Haman 1226 (holotype: GH!; isotype: US!). Encelia deltoidea Rusby, Desc. S. Amer. pl. 154. 1920.-TYPE: COLOMBIA. Magdalena: Playa Brava, seashore, on dry rocks and occasionally in sand, Nov 1898, Smith 594 (lectotype, here designated: NY!; isolectotypes: G! GH! S!). Simsia caucana Cuatrecasas, Revista Acad. Colomb. Ci. Exact. 9: 244. 1954. TYPE: COLOMBIA. Cauca: between Popayain and Purace, bridge over Anam bio River, 2500 m, 10Dec 1939, Perez & Cuatrecasas 5874 (holotype: US!; isotypes: COL! F!).

Suffrutescent, taprooted perennials to shrubs. Stems 0.5-4 m tall, 1-3 cm in diameter, erect, green to yellow, brown, or purple, canescent to pubescent, hirsute, or subsericeous, sometimes glandular. Leaves with petioles 0.6-5 cm long, 0.5-2 mm in diameter, at base with stipules of opposite leaves fused to form nodal discs 0-1.5 cm long and wide; blades 3-14.5 cm long, 1-13 cm wide, ovate to deltate, herbaceous, unlobed to deeply 3-5-lobed, base cordate to truncate or cuneate, apex acute to acuminate, pilose to densely sericeous, indument usually denser abaxially, eglandular or with intermixed short-stalked glands, margins crenate to dentate. Inflorescence corymbiform, moderately clustered; peduncles very short to 4.5 cm long, 0.5-2 mm in diameter, hirsute and intermixed glandular-puberulent and pubescent. Heads radiate, 10-16 mm long, 20-45 mm in diameter. Involucre campanulate, 7-14 mm long, 7-13 mm in diameter; phyllaries 16-35, 3-seriate, unequal, appressed to slightly reflexed apically, acute to acuminate at the tips, light green to greenish black, scabrous to sericeous, rarely glandular, margins entire; outer phyllaries 3.2-6 mm long, 1.4-2.8 mm wide, ovate to lance-ovate; inner phyllaries 7.8-14.4 mm long, 1.6-3.8 mm wide, lance-linear to lance-obovate. Pales 7.1-10.6 mm long, 1.2-3 mm wide, stramineous to purplish or speckled greenish black, acute to acuminate, antrorsely pubescent apically. Ray florets 8-19; ligules 5.5-14.5 mm long, 1.3-6 mm wide, light orange-yellow; tube 1.1-2.3 mm long, 0.4-0.6 mm in diameter, ovary 4.5-8.5 mm long, 0.4-0.5 mm wide. Disc florets 13-59; corolla 5.5-9 mm long, 1-1.5 mm in diameter; tube 0.8-2.3 mm long, 0.5-0.8 mm in diameter; lobes 0.6-1.5 mm long, 0.5-0.8 mm wide; anthers 2.9-4.5 mm long, yellow to black, terminal appendages stramineous; achenes 4-6.6 mm long, 1.7-3 mm wide, laterally flattened, awns 0.8-5 cm long, rarely absent, squamellae absent. Chromosome number: n = 17. Phenology. Flowering throughout the year. Distribution (Fig. 14). Throughout theAndes in northern South America from eastern Venezuela to northern Ecuador. Growing in areas of salt spray on beaches, in dry river valleys, and in cool, moist upland cloud forests, this species exhibits the greatest range of habitat variation in the genus. It is often found in highly disturbed 32 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 30

800 700 600 5Q0 400

FIGD|. 14. Distributio of Simsi dmb-.frtculoa, and theSouthAi...... can populations f0

S. foetida var. panamensis.

sites such as roadsides and old fields as well as in relatively undisturbed sites; 0 2600 m.

REPRESENTATIVE SPECIMENS: Venezuela. ME~RIDA: near La Carbonera, Aristeguieta 2476 (F, NY, US); outskirts of Chiguara, Badillo 5181 (MY); outskirts of Estainquez, Badillo 6420 (MY), 7167 (MY); Chama Valley between Ejido & Las Gonzales, Breteler 4545 (MO, NY, US); vicinity of Crist6bal Col6n, Broadway 463 (GH, NY, US); Lagunillas, Jahn 682 (US); D. Cpo. Elias, Gonzalez 614 (MY), Gonzalez et al. 215 (MER); between Ejido & Lagunillas, Oberwinkler 14120 (GZU, M); Brecenio Hacienda, Reed 596 (US).-MONAGAS:Via Guanaguana, Cumara 2146 (MY); Cerro Guachairo cave, near Guacharo, Steyermark 62345 (F, GH, MO, NY, US).-SUCRE: between Cumanocoa & Cocollar, Steyermark 62424 (F, K, NY, US). Colombia. BOYACA:Soata, Cuatrecasas & Barriga 1019 (COL, F); Boavita, Cuatrecasas & Barriga 1933 (COL, F, US).-CAUCA: Boquer6n del Dagua, Lehmann 4710 (K); bridge over Rio Anambio on W side of Purace, Spooner & Guevara 2933 (COL, 05); 2 kmW of bridge over 1990 SIMSIA 33

Rio Anambio on W side of Purace, Spooner & Guevara 2935 (COL, OS, WIS).-CUNDINAMARCA: Hacienda El Cucharo between Tocaima & Pubenza, Killip et al. 38378 (COL, F, US); 10 km SE of Chipaque, King & Guevara 5817 (COL, SMU, US); 46 km NW of Bogota, Olsen & Escobar 600 (LL, WIS); between La Mesa & Apulo, Spooner & Guevara 2921 (B, COL, OS).-HUILA: 11 km NW of Gigante, Davidse et al. 5802 (COL,MO), Spooner & Guevara 2924 (COL, OS, UC); 8 km S of Hobo, Spooner & Guevara 2923 (COL, OS, UC, US, WIS); 13 km fromNieva-Gigante Rd on rd to Tesalia, Spooner & Guevara 2925 (COL, OS, UC); 12 kmW of Paicol, Spooner & Guevara 2927 (BM, COL, OS).-MAGDALENA:Playa Brava, Burbidge 75/369 (NY); Playa Santa Maria, 26 Dec 1948 Reichel Dolmatoff s.n., (COL, US); beach at Santa Marta, Spooner 2975 (COL, OS, P, UC, US, WIS). NARINO: 80 km N of Pasto on rd to Popayan, Spooner & Guevara 2937 (COL, OS, P); 62 km N of Pasto on rd to Popaydn, 15 km N of JuanambuRiver crossing, Spooner & Guevara 3938 (COL, OS, OSH); 1 km N of N edge of Pasto on Rd to Popayan, Spooner & Guevara 2941 (COL, OS, WIS); by Rio Guaitara bridge on Pasto-Tulcin Rd, 1 km N of turnoff toTuquerres, Spooner & Guevara 2944 (COL, OS, W); 6 kmW of Pan-American Hwy on rd to Tuiquerres, Spooner & Guevara 2945 (COL, OS).-SANTANDER: 14 km S of Zapatoca toward La Fuente, Gentry & Forero 15458 A (COL, MO); Rio Surata valley between El Jaboncillo & Surata, Killip & Smith 16454 (F, GH, US).-TOLIMA: Chicoral, Haught 6374 (COL, NY, UC); 16 km W of Ibague on rd to Cajamarca, Spooner & Guevara 2972 (COL, OS); 26 km E of Ibague on rd to Gualanday, Spooner & Guevara 2973 (COL, OS).-VALLE: Rio Dagua, Lehmann 2964 (BM, G, GH, K, US); Yumbo, Pennell 5855 (GH, NY, PH, US), Spooner & Guevara 2952 (COL, GH, OS); 7 km NW of Cali on rd to Dagua, Spooner & Guevara 2953 (COL, OS); 1 km SE of S end of Dagua on rd to Cali, Spooner & Guevara 2956 (COL, OS); Rio Dagua, 4 km W of Loboguerrero, Spooner & Guevara 2957 (COL, OS, TEX). Ecuador. IMBABURA:12 km N of Ibarraon Rd toTulcain, Olsen & Escobar 496 (LL, NY, WIS), 497 (LL,WIS), Spooner 2909 (BM, LE, MEXU, OS, QCA, WIS); Salinas-Lita Rd., Panero & G6mez 1099 (TENN); S end of Ibarra,Spooner 2916 (GH, MO, OS, OSH, QCA, WIS).-PICHINCHA: Quito-Otavalo Rd near the town of Guayallabamba, Spooner 2918 (OS, TENN, QCA, UC, US, WIS).

Simsia fruticulosa is distinguished by its unequal phyllaries, sericeous leaves, and achenes larger (4-6.6 mm long) than those of the similar S. ghiesbreghtii (achenes 2.6-3.5 mm long). Simsia fruticulosa also is similar to S. molinae; S. fruticulosa has leaves pilose to densely sericeous and S. molinae has leaves short pilose or sometimes hispid. The heads of S. fruticulosa also tend to be borne on peduncles shorter than those of S. molinae. Simsia fruticulosa is highly variable. Seven species have been described based on differences of stem shape; leaf size, shape, and pubescence; stipule morphology; head size, shape, and pubescence; and achene pubescence. Some collections are quite distinctive, such as Spooner & Guevera 2933, 2935, 2936 with narrower leaves and longer outer phyllaries, or Spooner & Guevara 2923, 2973; King & Guevara 5817; Haught 6374; Breteler 4545 with larger heads and increased numbers of phyllaries, rays, and disc florets, but these and other variants were found scattered throughout the range of the species and all appear to be connected by intermedi ates. These observations indicate that all populations should be recognized as be longing to one highly polymorphic species. Triana's citation for S. pastoensis is "crescit prope Ortega, 1200 m, in prov. Pasto." Trianas's types are at COL, P, and other institutions (Stafleu & Cowan 1986). No specimens with these exact data have been located there or elsewhere. The best matches are two specimens from P: Nouvelle-Grenade, Prov. Pasto, Ortega, 1500 m, 1851-1857, Triana 1362, and Jun 1853, Triana 2553. The former is chosen as the lectotype. The exact locality of these collections is unknown, because Ortega is in the Province of Tolima. Dugand (1944) indicated that Triana's collec tions from "Prov. of Pasto" were made on the eastern side of the Andes in the Province of Narinlo. 34 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 30

The type of Encelia deltoidea, Smith 594, was collected inNovember, 1898. Many other collections labeled Smith 594 were made at the type locality but col lected on 26 Oct 1898, or the printed label has 1898-1901 and no other date added. These specimens are at CM, F, G, LL, MICH, MO, NY, UC, and US (5), and are regarded here as topotypes.

4. Simsia molinae H. Robinson & R. D. Brettell, Phytologia 24: 375. 1972.-TYPE: NICARAGUA.Estelfi: common along Esteli River, vicinity of Guava, 20 km [N] of Esteli, 5 Nov 1968,Molina 23122 (holotype: US!; isotypes: CAS! F! G! MO! NY!).

Suffrutescent, taprooted perennials to shrubs. Stems 1-3.5 m tall, 1-3 cm in diameter, erect, green to yellow or purple, puberulent to pubescent, weakly hispid. Leaves with petioles 1-4 cm long, 1mm in diameter, at base with stipules of opposite leaves fused to form nodal discs 0-2 cm long and wide; blades 3.5-11 cm long, 2.5-14 cm wide, ovate to deltate, herbaceous, unlobed to deeply 3-lobed, base cordate to truncate or cuneate, apex acute to acuminate, glandular-puberulent to short-pilose, sometimes with intermixed hispid hairs, margins crenate-dentate. Inflorescence corymbiform, loosely branched, heads subsessile or borne on peduncles up to 6 cm long, 0.5-1.5 mm in diameter, glandular-puberulent with intermixed short pubescent hairs. Heads radiate, 13-17 mm long, 20-45 mm in diameter. Involucre campanulate, 10-13 mm long, 8-15 mm in diameter; phyllaries 18-37, 3-seriate, unequal, appressed except slightly reflexed apically, acute to acuminate at the tips, yellow-green to brown, glandular-puberulent and short-pubescent, margins entire; outer phyllaries 3-8.3 mm long, 1-2.9 mm wide, ovate to ovate-lanceolate; inner phyllaries 9.5-14 mm long, 1-1.9 mm wide, lanceolate. Pales 8-11.1 mm long, 2-3 mm wide, stramineous, sometimes flecked purple or green-black at the apex, acute or acuminate, antrorsely pubescent apically. Ray florets 8-24; ligules 10-19.5 mm long, 3-6.5 mm wide, light orange-yellow; tube 1-2 mm long, 0.5-0.6 mm in diame ter; ovary 4.2-8 mm long, 0.5-0.7 mm wide. Disc florets 29-85; corolla 6-8 mm long, 1-1.5 mm wide; tube 1-1.8 mm long, 0.4-0.7 mm in diameter; lobes 1-1.5 mm long, 0.5-0.8 mm wide; anthers 3.9-4.5 mm long, yellow to black, terminal append ages stramineous; achenes 4.2-6.1 mm long, 2.5-3.5 mm wide, laterally flattened, awns 3.5-5.5 mm long, squamellae absent. Chromosome number: n = 17. Phenology. Flowering fromOctober throughNovember. Distribution (Fig. 12). Honduras to northern Costa Rica; moist thickets, road sides, and along streams; 0-950 m.

ADDITIONAL SPECIMENS EXAMINED. Honduras. MORAZAN: base of Cerro Majicaran, SE of El Zamorano, Molina 1413 (F, GH, MO), Standley 14009 (F, GH, US). Nicaragua. ESTELf: 5 km from Esteli, Molina 23020 (BM, CAS, S), 23027 (F, GH, U); Pan-American Hwy, 10.9 mi N of bridge over Esteli River on N side of Esteli, Spooner & Dorado 2701 (F, LE, MEXU, OS, OSH, TENN, TEX, UC, US, WIS); 20 km N of Esteli, Williams & Molina 42327 (MICH, US).-MANAGUA: Sierra de Managua, 1930-1940, Garnier s.n. (F); near El Crucero, along Hwy 8, 2.4 mi SW of Hwy 2, Spooner & Dorado 2704 (OS), Stevens 5171 (MO, OS); both sides of Pan-American Hwy, 2.7 mi SE of jct with Rt 8 to Masachapa, Spooner & Dorado 2714 (CR, F, MEXU, MO, OS, OSH, P, TEX, UC, US, WIS). Costa Rica. GUANACASTE: Playas del Coco, Cicci6 45 (CR), Spooner 2901 (CR, OS).

Simsia molinae is similar to S. fruticulosa in its habit, nodal structure, leaf shape, phyllaries, color, and variability of anther color. Simsia molinae 1990 SIMSIA 35

differs mainly in its sparser leaf pubescence and themore open inflorescence (see discussion under S. fruticulosa). Although one might consider treating this taxon as a variety of S. fruticulosa, its relative ease of recognition and allopatric range more strongly suggest specific status.

5. Simsia holwayi S. F. Blake, Contr. Gray Herb. 52: 46. 1917.-TYPE: GUATE MALA.Guatemala: Agua Caliente, on Barrios-Guatemala City Railway, 4 Feb 1917, Holway 854 (holotype: GH!; isotype:US!)

Suffrutescent, taprooted perennials to shrubs. Stems 0.5-4 m tall, 0.3-1 cm in diameter, green to purple to brown, hirsute and pubescent and intermixed glandular-puberulent. Leaves with petioles 1-4 cm long, 0.5-1 mm in diameter, at base with stipules of opposite leaves fused to form nodal discs 0-1 cm high and wide; blades 3-8 cm long, 2-6 cm wide, ovate to deltate, herbaceous, unlobed to 3 lobed one-half way to midrib, base cordate to truncate to cuneate, apex acute or acuminate, glandular, short-hispid and pilose adaxially, longer pilose abaxially, margins crenate to dentate. Inflorescence corymbiform, loosely branched; heads subsessile or borne on peduncles up to 8 cm long, 0.5-1 mm in diameter, glandular puberulent and short-hirsute. Heads radiate, 10-13 mm long, 10-15 mm in diame ter. Involucre campanulate to ovoid, 9-11 mm long, 6-7 mm in diameter; phyllaries 18-27, 3-seriate, unequal, appressed, acute to acuminate at the tips, usually purple and green, puberulent, sometimes glandular and pubescent, margins entire; outer phyllaries 3-4.5 mm long, 1.5-2.2 mm wide, ovate; inner phyllaries 7.2-9.5 mm long, 1.2-1.8 mm wide, lanceolate to linear. Pales 8-9.2 mm long, 1.7 2.8 mm wide, stramineous, purple or green-black apically, acute to acuminate, antrorsely pubescent apically. Ray florets 8-9, ligules 5-7 mm long, 2.5-2.9 mm wide, light orange-yellow; tube 0.8-1 mm long, 0.3 mm in diameter; ovary 6-7.5 mm long, 0.5 mm wide. Disc florets 20-31; corolla 5.7-6.7 mm long, 1-1.6 mm in diameter; tube 0.9-1.2 mm long, 0.4-0.7 mm in diameter; lobes 0.7-1.1 mm long, 0.6-0.8 mm wide; anthers 3.5-3.8 mm long, yellow, terminal appendages strami neous; achenes 4.7-5.6 mm long, 2.3-3 mm wide, laterally flattened, awns 3.6-4.2 mm long, squamellae absent. Chromosome number: n = 17. Phenology. Flowering fromOctober throughFebruary. Distribution (Fig. 12). Guatemala; along the Monagas Valley and adjacent slopes; in dry tropical deciduous forests; 600-900 m.

ADDITIONAL SPECIMENS EXAMINED: Guatemala. EL PROGRESO: on rd to Coban from Guatemala Puerto Barrios Rd, near Morazan, 7.5 mi S of Baja Verapaz-El Progreso border, Spooner & Dorado 2746 (BM, COL, MEXU, NY, OS, TEX, UC, WIS); Guatemala-Puerto Barrios Rd, 2.4 mi SW of rd to Sanarate, Spooner & Linder 2750 (MEXU, OS); Guatemala-Puerto Barrios Rd, 0.9 mi NE of rd to Sanarate, Spooner & Linder 2759 (MEXU, OS).-GUATEMALA: on railroad cut, 0.2 mi NE of railroad station at Agua Caliente, Spooner & Dorado 2692 (MEXU, OS, OSH, P, TENN, UC, US, WIS).

Simsia holwayi is distinguished by its nodal discs, unequal phyllaries, yellow anthers, short ray corollas (5-7 mm long), and glandular, hispid, and pilose leaves. The purple phyllaries and campanulate-ovoid heads with slightly constricted apices give it a superficial resemblance to S. lagascaeformis, which is an annual species without nodal discs. 36 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 30

All populations of this species are found along the dryMonagas River Valley and adjacent slopes inGuatemala. A specimen (Holway 854) with similar appearance and identical label data except "10 Feb 1917" is deposited at US. The date is regarded as a typographical error, and the specimen is considered an isotype.

6. Simsia steyermarkii H. Robinson & R. D. Brettell, Phytologia 24: 375. 1972. TYPE:GUATEMALA. Zacapa: Sierra de las Minas, trail between Santa Rosalia de Marmol and Vegas, 19 Jan 1942, Steyermark 42931 (holotype: US!; isotype: F!).

Suffrutescent, taprooted perennials to shrubs. Stems 1-4 m tall, 0.5-3 cm in diameter, erect, green to purple or brown, puberulent andmildly scabrous. Leaves with petioles 1-7 cm long, 1mm indiameter, at base with stipules of opposite leaves fused to form nodal discs 0-2 cm long and wide; blades 3-15 cm long, 2-9 cm wide, ovate to deltate, herbaceous, unlobed, base cordate to truncate or cuneate, apex acute to acuminate, short-hispid and puberulent, sometimes glandular adaxially, glandular-puberulent abaxially, margins subentire to crenate. Inflorescence corym biform, usually tightly clustered; heads subsessile or borne on peduncles up to 2 cm long, 1mm indiameter, glandular-puberulent and hispid. Heads radiate, 11-13 mm long, 13-17 mm in diameter. Involucre campanulate, 10-12 mm long, 6-11 mm in diameter; phyllaries 18-21, 3-seriate, unequal, appressed except slightly reflexed apically, acute to acuminate at the tips, green to green-black to black, scabrous, sometimes puberulent and glandular, margins entire; outer phyllaries 2-5.9 mm long, 0.7-2 mm wide, ovate to lance-ovate; inner phyllaries 7.5-9.7 mm long, 1.3 1.7 mm wide, lanceolate. Pales 7.8-9.5 mm long, 2.2-2.7 mm wide, stramineous, sometimes speckled green-black apically, acute to acuminate, antrorsely pubescent apically. Ray florets 8; ligules 8-10 mm long, 3.5-4.5 mm wide, light lemon-yellow; tube 1.2-1.6 mm long, 0.3-0.5 mm in diameter; ovary 4.3-6.3 mm long, 0.4-0.6 mm wide. Disc florets 15-34; corolla 5.8-7.2 mm long, 1.2-1.5 mm in diameter; tube 0.8-1 mm long, 0.5-0.7 mm in diameter; lobes 0.7-1.2 mm long, 0.6-1.1 mm wide; anthers 3.5-4.2 mm long, yellow to black, terminal appendages stramineous; achenes 3.2-4.2 mm long, 1.8-2.5 mm wide, laterally flattened, awns absent or 1.9 2.7 mm long, squamellae absent. Chromosome number: n = 17. Phenology. Flowering fromNovember through January. Distribution (Fig. 12). Guatemala, in the Sierra de lasMinas and adjacent uplands to the west; in pine or pine-oak forests; 1300-1500 m.

ADDITIONAL SPECIMENS EXAMINED: Guatemala. ALTA VERAPAZ: rd to Sacapulas, 5 km WNW of San Crist6bal Verapaz, King 7337 (field specimens: MO, NY; greenhouse specimens from seed: B, CAS, G, M, NY, U, US, WIS).-BAJA VERAPAZ: 4.8 mi from San Miguel Chicaj town square on rd to Cubulco, Spooner & Dorado 2748 (BM, GH, MEXU, MICH, OS, OSH, TENN, TEX, UC, US).

Simsia steyermarkii is most easily distinguished by its aggregated inflores cences, small achenes (3.2-4.2 mm long), and light lemon-yellow ligules.All speci mens examined, including greenhouse-grown specimens of Spooner & Dorado 2748, have aggregated inflorescences; greenhouse-grown specimens from of King 7337 have more open inflorescences, but are otherwise typical of the species. 1990 SIMSIA 37

7. Simsia annectens S. F. Blake, Contr. Gray Herb. 52: 43. 1917.-TYPE: MEXICO. Mexico: Zuetepec, Almoloya de lasAlguisiras, in bushes along edge of rd, 23 Dec 1904, Seler & Seler 4472 (holotype: GH!; isotypes: ENCB! GH!).

Suffrutescent, taprooted perennials to shrubs. Stems 1.5-4 m tall, 1-3 cm in diameter, erect, green to brown or purple, usually with harsh, hispid hairs, some timesmerely hirsute or pubescent, sometimes glandular. Leaves with petioles 1-14 cm long, 1-2 mm in diameter, often winged throughoutwith wings to 1.5 cm wide, or partlywinged by the decurrent blade, or without wings, at base perfoliate or with stipules of opposite leaves fused to form nodal discs 0-5.5 cm long andwide; blades 3-25 cm long, 2-30 cm wide, ovate to deltate, herbaceous, unlobed to deeply 3-5 lobed, base cordate to truncate or cuneate, apex acute or acuminate, usually sca brous and hirsute, especially adaxially, sometimes glandular-puberulent, especially abaxially, margins crenate to dentate. Inflorescence corymbiform, of tightly clus tered heads or more lax; peduncles 0.5-7 cm long, 1-2 mm in diameter, sometimes widely divaricately branched, usually hirsute, sometimes intermixed glandular puberulent and pubescent. Heads radiate, 11-18 mm long, 15-30 mm in diameter. Involucre campanulate, 9-12 mm long, 6-15 mm in diameter; phyllaries 14-29, 2 3-seriate, unequal to subequal, appressed to greatly reflexed, green to yellow green, dark blackish green or rarely tingedwith purple, short-puberulent and sca brous to long-hispid, sometimes glandular, acute to long-attenuate, margins entire; outer phyllaries 3.7-11 mm long, 1.4-2.8 mm wide, broad-ovate to lanceolate; inner phyllaries 7.5-15 mm long, 1.5-2.5 mm wide, lanceolate to linear-obovate, rarely sinuate. Pales 7.2-10 mm long, 1.7-3.3 mm wide, stramineous, acute to acuminate, subglabrous or antrorsely pubescent apically. Ray florets 8-14; ligules 6.5-14 mm long, 2-7 mm wide, light lemon-yellow; tube 0.8-2.2 mm long, 0.3-0.7 mm in diameter; ovary 3.7-8.2 mm long, 0.5-1 mm wide. Disc florets 21-67; corolla 5.5-7.1 mm long, 1-1.7 mm in diameter; tube 0.7-2.1 mm long, 0.4-0.8 mm in diameter; lobes 0.6-1.2 mm long, 0.5-1 mm wide; anthers 3.2-4.4 mm long, yellow to black, terminal appendages stramineous; achenes 3.5-7 mm long, 1.7-3.6 mm wide, laterally flattened, awns absent or 1.9-3.6 mm long, squamellae absent. Chromosome number: n = 17. Simsia annectens usually is easily distinguished by distinctive perfoliate leaves or conspicuously winged petioles, but some nodes or some plants in a population may exhibit unwinged petioles; in such cases prominent nodal discs connecting the bases of adjacent leaves are usually present. Simsia annectens exhibits much variation in pubescence of all parts, leaf shape, inflorescence branching patterns, head size, and phyllary gradation and posture. Stem pubescence varies from hirsute to so harshly hispid that individual trichomes can even penetrate skin. Leaf pubescence varies from eglandular to glandular, hispid to puberulent. As in all species of Simsia with lobed leaves, entire leaves also are found, sometimes even on plants with mostly lobed leaves. Petioles vary in size and alation from completely unwinged to winged throughout with a width up to 1.5 cm, then perfoliate. The inflorescence often is a tight cluster of many heads, with the peduncles widely divergent. Although quite distinctive in some populations (e.g., Hinton 12285, Mexia 1832), this inflorescence pattern is far from a constant feature and is not useful taxonomically. Phyllaries are gradate, appressed, and 38 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 30

scabrous, or subequal, reflexed, and long-setose. The size of the heads is variable and not taxonomically diagnostic in this species. The conspecific nature of all populations here grouped under S. annectenswas recognized only after observations of intra- and interpopulational variation from throughout the range of the species. Two distinct but occasionally intergrading varieties are here recognized: var. annectens, with densely setose and greatly re flexed phyllaries, and var. grayi with short-puberulent, sometimes scabrous phyl laries, which are appressed or slightly reflexed apically. Occasional intermediates connect these extremes (e.g., Hinton 2693, Matuda 26954, Spooner & Villasefior 2566).

7a. Simsia annectens var. annectens. Simsia hintonii H. Robinson & R. D. Brettell, Phytologia 24: 368. 1972. TYPE:MEXICO. : Mina, Puerto Rico, 1800m, by forest stream, 10 Dec 1939, Hinton et al. 14976 (holotype: US!; isotypes: GH! K! LL! MICH! UC!). Simsia subsetosa H. Robinson & R. D. Brettell, Phytologia 24: 376. 1972. TYPE: MEXICO. Mexico: , Rincon del Carmen, 1340 m, woods, 23 Nov 1932, Hinton 2693 (holotype: US!, photo: US!; isotypes: BM! DS! G! MICH! NY!).

Plants 1.5-4 m tall. Phyllaries usually greatly reflexed, long-hispid, attenuate apically; outer phyllaries 5.2-11.5 mm long, 1.4-2.5 mm wide, lance-ovate; inner phyllaries 7-15 mm long, 1.6-2.5 mm wide, lanceolate. Chromosome number: n = 17. Phenology. Flowering from September throughMarch. Distribution (Fig. 15). Mexico, along the Transverse Volcanic Belt in Jalisco, Michoacain, and Mexico, and the Sierra Madre del Sur inMichoacan and Guerrero; in open pine-oak forests, on mountain slopes, and along streamsides; 1000-2400 m.

ADDITIONAL SPECIMENS EXAMINED. Mexico. GUERRERO: 36 km NE of Puerto del Gallo along rd to Filo de Caballo, Breedlove 36152 (CAS, MICH, MO), Spooner 2845 (MEXU, MICH, OS, TENN, TEX, UC, US, WIS); Chichihualco, Cruz de Ocote, 25 kmWSW of Camotla, Feddema 2866 (MICH), Rzedowski 18109 (ARIZ, ENCB, MICH); Mina, Manch6n, Hinton 11271 (GH, K, LL, MICH, NY, UC, US, W); Montes de Oca, Vallecitos, Hinton 11651 (GH, K, LL, MICH, NY, UC, US); Mina, Campo Morado-Otatlan, Hinton 14843 (GH, K, LL, MICH, NY, UC, US); rd to Chilpancingo, 16 km E of Aserradero Agua Fria, Rzedowski & McVaugh 272 (ENCB, MICH); 3-5 mi W of Rt 95 on rd to Chilpancingo, Stuessy & Gardner 4225 (ASU, ENCB, MO, OS, WIS).-JALISCO:Huejotitan, Oct 1912, Diguet s.n. (F, MICH); Hacienda del Ototal, Arroyo del Ototal, W of San Sebastian, Mexia 1832 (BM, CAS, DS, F, G, GH, MICH, MO, NY, UC).-MfxIco: Temascaltepec, Temascaltepec, Hinton 2801 (BM, G, K), Matuda 30468 (ENCB); Temascaltepec, Ixtapan, Hinton 2935 (BM, G, MEXU, MO); Tejupilco, Ocotepec, Rzedowski 25298 (ENCB).-MICHOACAN:Coalcoman, Pto. Zarzamora, Hinton 12714 (K, LL, MICH, NY, S, UC); Coalcoman, Coalcoman, Hinton 12885 (ARIZ, CHAPA, ENCB, GH, LL, MO, NY, PH, US, W); along Rt 15, 16.7 mi E of E limits of Morelia between Rio El Salto & village of Polvilla, Stuessy & Gardner 3108 (ASU, F, MEXU, MICH, NY, OS, UC). Intermediates between var. annectens and var. grayi: MexIco: rd between Valle de Bravo & Colorines, Matuda 26954 (MEXU, NY), Spooner & Villasefior 2566 (GH, LE, MEXU, OS, TEX, UC, US, WIS), Villasefior & Spooner 766 (MEXU, OS).

Many collections of S. annectens var. annectens are densely glandular and have a yellow-green coloration similar to that of S. foetida. Some collections (e.g., 1990 SIMSIA 39

1020 960 900

*0 200 200 AAA

S. annectens * var. annectens A var. grayi

x S. villasenorii

960 90o

FIG. 15. Distribution of Simsia annectens var. annectens, S. annectens var. grayi, S. tenuis, and S. villasenorii.

Spooner 2845) even have sinuate phyllaries like those of S. foetida, but the spicy odor of S. foetida was absent from this collection. This combination of greenish yellow, densely glandular leaves and sinuate phyllaries suggests the possibility of earlier hybridization of the two species. Robinson and Brettell (1972) placed much taxonomic weight on the widely divergent branching pattern of the inflorescence of S. annectens, but this pattern is far from constant and of limited taxonomic utility. A focus on this feature possibly led them to recognize S. subsetosa. The type of S. subsetosa differs from many other populations of S. annectens by its sparse pubescence, but similar sparse pubescence is present in other populations of both varieties (e.g., var. annectens:Hinton 1285; var. grayi: Breedlove & Strother 47040, Spooner 2788). Simsia hintonii was distinguished from S. annectens by the reduced alation of the petioles, more hirsute phyllaries, puberulous rather than densely scabrous leaf pubescence, and less glandular stems. All of these differences are well within the range of variation present in S. annectens var. annectens.

7b. Simsia annectens var. grayi (Schultz-Bipontinus ex S. F. Blake) D. M. Spooner, comb. et stat. nov. Simsia grayi Schultz-Bipontinus ex S. F. Blake, J. Wash. Acad. Sci. 18: 26. 1928.-TYPE: MEXICO. Oaxaca, Liebmann 561 (holotype: P!, fragment: US!, photos: OS! US!; isotypes: C[2]!, photo and fragment:US!, tracing:GH!).

Plants 1.5-3 m tall. Phyllaries appressed to only slightly reflexed at the very apex, short-puberulent to scabrous, acute to acuminate apically; outer phyllaries 3.7-6 mm long, 1.6-2.8 mm wide, ovate to lance-ovate; inner phyllaries 7.5-9.1 40 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 30 mm long, 1.5-2.4 mm wide, lanceolate to linear or linear-obovate. Chromosome number: n = 17. Phenology. Flowering from September throughNovember. Distribution (Fig. 15). Mexico, along the Transverse Volcanic Belt, in the SierraMadre del Sur of Guerrero, and along the northern highlands of Chiapas and Oaxaca; in openings in pine-oak forests, often in shallow soil of volcanic and limestone rocks; 850-1930 m.

ADDITIONAL SPECIMENS EXAMINED. Mexico. CHIAPAS: N of Tuxtla Gutierrez at mirador for Chicoasan Dam, Breedlove & Strother 47040 (CAS), Spooner 2788 (LE, MEXU, MICH, OS, OSH, TENN, TEX, UC, US, WIS).-GUERRERO: Zihuatenejo-Cd. Altamirano Rd, 81 km N of the Coast Hwy, 3 km N of highest pass on hwy, Barrie et al. 650 (TEX).-MICHOACAN: 7 mi SE of Cd. , Cronquist 10314 (CAS, DUKE, ENCB, F, GH, MEXU, MICH, NY, TEX, US); 2 mi E of Tuxpan, Crutchfield & Johnston 5902 D (TEX); Zitacuaro-San Jose Purua, Hinton 13260 (B, ENCB, F, G, LL, MICH, NY, S, UC, W); along dirt rd to Rinc6n de la Dolores, 4 km SW of Cd. Hidalgo, Iltis et al. 710 (OS, WIS); Rt 15, 1.8 mi S of Tuxpan town square, Keil & Canne 9081 (NY, OS), Spooner & Brandenburg 2279 (CHAPA, MEXU, NY, OS); Rt 15, 14 mi W of Cd. Hidalgo, Melchert et al. 6428 (ENCB, MICH, TEX); rd to Jungapeo, 1mi S of rd to ZitAcuaro, Panero & Schilling 443 (TENN); along Hwy 15, 0.2 mi W of rd toMaravatio, Spooner & Villasefior 2562 (COL,MEXU, MICH, LE, OS, OSH, UC, US, WIS), Villasefior & Spooner 762 (MEXU, OS); along Hwy 15, 2.5 mi E of jct of rd to Maravatfo, Spooner & Villasenior 2563 (B, GH, MEXU, MO, NY, OS, TEX, UC, US, WIS); 3mi E of Tuxpan, Spooner & Villasefior 2564 (MEXU, OS, US).-OAXACA: Oaxaca, Buchinger 497 (P); Tepinapa, Liebmann 560 (US).

Itwas my initial impression that the populations of S. annectens var. grayi from Guerrero and Michoacain were taxonomically distinct from the populations from Oaxaca and Chiapas. The only complete population samples showing populational variability and the full development of the perfoliate leaves are the recent collec tions from Chiapas. These southern populations tend to have less strigose stems, more densely glandular leaves, and less widely divergent branches of the inflore scence. Further study indicated considerable intergradationof these features in the north.

8. Simsia tenuis (Fernald) S. F. Blake, Proc. Amer. Acad. Arts 49: 380. 1913. Encelia tenuisFernald, Proc. Amer. Acad. Arts 33: 94. 1897.-TYPE: MEX ico. Guerrero: Acapulco and vicinity, Nov 1896 (fideMcVaugh, 1956), Palmer 96 (holotype: GH!; isotypes: F[2]! K! MO[2]! NY! UC! US!).

Taprooted annuals to suffrutescent perennials. Stems 0.5-1.5 m tall, 0.1-1.1 cm in diameter, erect, green to yellow or purple, subglabrous toweakly hirsute and pubescent and intermixed glandular-puberulent. Leaves with petioles 0.5-8 cm long, 0.5-1 mm in diameter, at base with stipules of opposite leaves fused to form nodal discs 0-1.2 mm long and wide; blades 2.5-13 cm long, 1.5-10 cm wide, ovate to deltate, herbaceous, usually unlobed, rarely 3-lobed, base cordate to truncate or cuneate, apex acute to acuminate, hirsute and intermixed glandular-puberulent, margins crenulate. Inflorescence corymbiform, loosely branched; peduncles 0.5-10 cm long, 0.5-1 mm in diameter, hirsute and intermixed glandular-puberulent and pubescent. Heads radiate, 9-14 mm long, 15-30 mm in diameter. Involucre campanulate, 6-11 mm long, 5-7 mm in diameter; phyllaries 12-25, 3-seriate, unequal, appressed except slightly reflexed apically, acute to acuminate at the tips, 1990 SIMSIA 41

light green to purple, scabrous and glandular, sometimes hirsute, margins entire; outer phyllaries 3.5-6 mm long, 1.5-2 mm wide, ovate to lance-ovate; inner phyllaries 6-11 mm long, 1-2 mm wide, lanceolate. Pales 5-8 mm long, 1.5-2 mm wide, stramineous, acute to acuminate, antrorsely pubescent apically. Ray florets 5-10; ligules 5.5-15 mm long, 1.5-7 mm wide, light orange-yellow; tube 1-2 mm long, 0.5 mm in diameter; ovary 4-7.5 mm long, 0.4-0.5 mm wide. Disc florets 12 14; corolla 7mm long, 1mm in diameter; tube 0.7-2 mm long, 0.5 mm indiameter; lobes 0.7-1.2 mm long, 0.5-0.8 mm wide; anthers 3.2-3.5 mm long, yellow to black, terminal appendages stramineous; achenes 3.2-6.1 mm long, 2-3.8 mm wide, laterally flattened, pappus absent. Chromosome number: n = 17. Phenology. Flowering fromAugust throughDecember. Distribution (Fig. 15).Mexico, in the SierraMadre del Sur near Acapulco, and in the lowlands of theRio Balsas Basin inMichoacain; in tropical deciduous forests; 300-600 m.

ADDITIONAL SPECIMENS EXAMINED. Mexico. GUERRERO: Acapulco, Sierra, Paray 2336 (ENCB). MICHOACAN:11-13 kmWSW of Apatzingan on rd toDos Aguas, Dieterle 4351 (ENCB, MICH, TEX); 5 km SW of Tepalcatepec on rd to Coalcoman, Spooner & Villasefior 2550 (B, CAS, CHAPA, COL, ENCB, F, FI, G, GA, GH, GUADA, K, LE, LL, MA, MEXU, MICH, NY, OS, OSH, P, SRSC, TENN, TEX, UC, US, VEN, WIS, XAL), Villasefior et al. 292 (MEXU), Villasefior & Spooner 742 (MEXU, OS).

This species is distinctive with its small heads and thin leaves. The two speci mens from Guerrero differ from those from Michoacain by possessing black vs. yellow anthers and disc florets with smaller tubes, but these features also vary in related species. The Guerrero and Michoacain populations otherwise are similar and are considered conspecific. Some of the individuals of Spooner & Villasenior 2550 were annuals, and others had suffruticose bases.

9. Simsia villasenorii D. M. Spooner, sp. nov.-TYPE: MEXICO. Chiapas: Mpio. of Cintalapa, crest of the Sierra with Pinus and Quercus and riparian situa tions with seasonal evergreen forest near the microwave station of La Mina, 12 km S of Mexican Hwy 190 near Rizo de Oro, 1000 m, 16 Oct 1971, Breedlove & Thorne 20576 (holotype DS!; isotypes: ENCB! LL! MEXU! MICH! MO! NY!).

Plantae lignosae, 0.5-2 m altae, petiolis foliorum 0.5-6 cm longis, stipulis foliorum oppositorum connatis, laminis ovatis vel deltatis, rigidis, non lobatis, 3-10 cm longis, 2-7 cm latis, basi cordatis vel truncatis vel cuneatis, apice acutis, involucris gradatis. Suffrutescent, taprooted perennials to shrubs. Stems 0.5-2 m tall, 0.3-3 cm in diameter, erect, green to purple, puberulent and scabrous. Leaves with petioles 0.5-6 cm long, 1-2 mm in diameter, at base with stipules of opposite leaves fused to form nodal discs 0-2 cm long and wide; blades 3-10 cm long, 2-7 cm wide, ovate to deltate, stiff, unlobed, base cordate to truncate or cuneate, apex acute, puberulent and scabrous, sometimes glandular, margins crenate-dentate. Inflorescence corym biform, tightly to more loosely branched; heads subsessile or borne on peduncles up to 7 cm long, 1 mm in diameter, glandular-puberulent and scabrous. Heads radiate, 15-18 mm long, 25-30 mm in diameter. Involucre campanulate, 14-15 mm 42 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 30

long, 8-11 mm in diameter; phyllaries 19-28, 3-seriate, unequal, appressed, acute to aristate at the tips, green to purple or brown, scabrous, sometimes glandular, margins entire; outer phyllaries 3.2-5.5 mm long, 1.2-2.3 mm wide, ovate; inner phyllaries 10-13 mm long, 1.5-2.1 mm wide, linear-lanceolate, linear to linear obovate. Pales 10-13.4 mm long, 2-2.6 mm wide, stramineous, purple apically, acute to acuminate, antrorsely pubescent apically. Ray florets 11-16; ligules 7.4-14 mm long, 2.5-4.5 mm wide, light lemon-yellow; tube 1.7-3.5 mm long, 0.3-0.5 mm in diameter; ovary 7.4-10.2 mm long, 0.5-1 mm wide. Disc florets 21-39; corollas 6.5-6.7 mm long, 1.5-2.1 mm in diameter; tube 0.7-1.1 mm long, 0.6-0.9 mm in diameter; lobes 1.1-1.3 mm long, 0.8-1.2 mm wide; anthers 3.5-4.1 mm long, yellow to black, terminal appendages stramineous; achenes 5.5-7.3 mm long, 2.5-3.5 mm wide, laterally flattened, awns 4.5-6.5 mm long, squamellae absent or 0.5-1 mm long. Chromosome number: n = 17. Fig. 16. Phenology. Flowering from September throughNovember. Distribution (Fig. 15). Mexico, inOaxaca in the SierraMadre del Sur, and in Chiapas in the Sierra Madre de Chiapas; in areas of tropical deciduous forest and oak-pine woods, including recently burned oak-pine woods; 800-1000 m.

ADDITIONAL SPECIMENS EXAMINED. Mexico. CHIAPAS: 10 km NW of Rizo de Oro along logging rd to Colonia Figar6a, Breedlove 44455 (DS); near microwave station of La Mina, Breedlove 52914 (CAS), 56333 (CAS), Breedlove & Almeda 60160 (CAS), Spooner 2789 (BM, MEXU, MO, OS, WIS). OAXACA: Pan-American Hwy, 93 mi NW of Tehuantepec & 59 mi SE of Oaxaca, Cronquist 9687 (GH, MEXU, MICH, NY, TEX); 57 mi SE of Oaxaca, Hartman & Funk 4169 (OS, US); Pan-American Hwy, 0.8 mi NW of jct of rd to San Carlos Yautepec in town of El Camar6n, Spooner 2816 (MEXU, OS).

Simsia villasenorii is a distinctive species. All parts of the plant are firm and stiff.Most species of Simsia have more herbaceous leaves; only S. santarosensis has leaves that are as firm.These differences aremost evident under wilting conditions in the greenhouse. Chiapan populations have an inconstant tendency to make tight clusters of axillary inflorescences. Oaxacan populations grow in dry, tropical deciduous for ests, the Chiapan in oak-pine woodlands. Spooner 2789 fromChiapas came from a burned woodland inwhich many shoots were growing from persistant rootstocks. Both the Oaxacan and Chiapan populations include some individualswith pappus squamellae between the awns. Such squamellae are uncommon in Simsia. This species is named in honor of Jose Luis Villasenior Rios, an authority on the Compositae of Mexico. He generously gave me one month of his time in the field, ledme tomany interesting localities of Simsia, and taughtme much aboutMexican Compositae.

10. Simsia setosa S. F. Blake, Proc. Amer. Acad. Arts 49: 379. 1913.-TYPE: MEXICO.: Alamos, 16-30 Sep 1890, Palmer 741 (holotype: GH!; isotypes: CAS! K! LE! MEXU! NY! UC! US[2]!).

Herbaceous perennials to suffrutescent, taprooted perennials. Stems 1-1.5 m tall, 0.3-1 cm in diameter, erect, green to yellow, brown, or purple, glandular puberulent, hispid and hirsute. Leaves with petioles 1-7.5 cm long, 1mm in diame ter, at base with stipules of opposite leaves fused to form nodal discs 0-2 cm long and wide; blades 3-9 cm long, 2-7 cm wide, ovate to deltate, herbaceous, unlobed to deeply 3-5-lobed, base cordate, truncate, or cuneate, apex acute to acuminate, 1990 SIMSIA 43

I~~~~~~~~~~~~IC

tA~~~~~ c

FIG. 16. Simsia villasenorii (Spooner 2789). A. Ray floret. B. Stem with mature inflorescence. C. Head. D. Achene. E. Pale. F. Disc floret. densely glandular-puberulent, hispid and hirsute, margins crenate to dentate. Inflorescence corymbiform, loosely branched; peduncles 1-7 cm long, 1 mm in diameter, glandular-puberulent and hirsute. Heads radiate, 13-18 mm long, 15-25 mm in diameter. Involucre campanulate, 12-15 mm long, 9-15 mm in diameter; phyllaries 14-28, 3-seriate, unequal, appressed except slightly reflexed apically, 44 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 30

83?

______

830

FIG. 17. Distribution of Simsia calva and S. setosa.

light green to brown-yellow, rarely lightly tingedwith purple, glandular-puberulent and hispid, margins entire; outer phyllaries 4-7 mm long, 1.5-2.2 mm wide, ovate to lance-ovate; inner phyllaries 11.5-13 mm long, 1.7-2.4 mm wide, lance-ovate. Pales 8.7-11 mm long, 2-2.8 mm wide, stramineous, sometimes speckled greenish black apically, acute to acuminate, antrorsely pubescent apically. Ray florets 8-13; ligules 7.5-9.5 mm long, 2.5-3 mm wide, light lemon-yellow; tube 1-1.2 mm long, 0.5 mm in diameter; ovary 7-10 mm long, 0.5-0.6 mm wide. Disc florets 18-40, corolla 6.6-8.5 mm long, 1-1.2 mm in diameter; tube 0.9-1.2 mm long, 0.5-0.7 mm in diameter; lobes 0.8-1.2 mm long, 0.7-0.9 mm wide; anthers 3.7-4.2 mm long, yellow to black, terminal appendages stramineous; achenes 5.2-6.5 mm long, 2.6-4 mm wide, laterally flattened, awns absent or 2.5-3.5 mm long, squamellae absent. Chromosome number: n = 17. Phenology. Flowering from July throughNovember. 1990 SIMSIA 45

Distribution (Fig. 17). Mexico, in the Sierre Madre Occidental in southern Sonora and adjacent and in Nayarit; in lowland tropical deciduous forests to higher elevation pine-oak forests, including recently burned-over pine woodlands; 400-1600 m.

ADDITIONAL SPECIMENS EXAMINED. Mexico. CHIHUAHUA: Guasaremos, Rio Mayo, Gentry 2464 (ARIZ, DES, F, GH, K, MO, S, UC); Rancho Beverly, Sierra Charuco, Langille 118 (DES), 173 (DES).-NAYARIT: along footpath from Jesus Maria to La Mesa, Colunga & Zizumbo 21 (MEXU), Spooner & Villasenior 2528 (BM, COL, GH, LE, MEXU, MO, NY, OS, OSH, P, TENN, TEX, UC, US, WIS).-SoNoRA: Sierra de Las Papas, Sonora-Chihuahua boundary, Gentry 654 (MICH); Arroyo Cuchuhaqui, Dist. Alamos, Gentry 822M (MICH); Cafi6n Sapopa, Rio Mayo, Gentry 1036 (DES, F, GH, MO, S); Cafi6n Estrella, Dist. Alamos, Gentry 3430 (CAS,MICH); Cafi6n de Bavispe, White 3120 (GH, MICH); Puerto de losAserraderos, Rancho de Cruz Diaz, White 3292 (MEXU, MICH); Cafi6n de la Palomita, N of El Tigre, White 4155 (GH, LL, MICH).

Simsia setosa is recognized by its densely glandular and hispid vesture, lobed leaves, and unwinged petioles. This species has a conspicuous yellowish appearance and spicy-smelling leaves similar to those found in S. foetida. This odor was not apparent in the field, but itwas evident in greenhouse-grown specimens raised from achenes of Spooner & Villasefior 2528.

11. Simsia sanguinea A. Gray, Smithsonian Contr. Knowl. 3 (5) (P1. wright. 1): 107. 1852. Encelia sanguinea (A. Gray) Hemsley, Biol. cent.-amer., Bot. 2: 185. 1881.-TYPE: MEXICO.[Probably Chiapas], mountains, temperate re gion, Ghiesbreght 305 (lectotype, designated by Blake, 1913:GH!). Simsia erythranthema Schultz-Bipontinus ex A. Gray, Proc. Amer. Acad. Arts 19: 9. 1883, pro syn. Encelia sanguinea var. palmeri A. Gray in S. Watson, Proc. Amer. Acad. Arts 22: 427. 1887. Simsia sanguinea var. palmeri (A. Gray) S. F. Blake, Proc. Amer. Acad. Arts 49: 395. 1913.-TYPE: MEXICO. Jalisco: Rio Blanco, Sep 1886, Palmer 602 (holotype: GH!; isotypes: BM! G! K! LE! MEXU! MO! NY [2]!US!). Helianthus hastatus Sesse & Mocinlo, P1. nov. Hisp. 147. 1890.-TYPE: MEXICO. Guerrero: "Chilapae" [=Chilapa] Sep [1789 fideMcVaugh, 1977], Sesse'et al. s.n. (holotype: MA; isotype: F!). Simsia triloba S. F. Blake, Proc. Amer. Acad. Arts 49: 393. 1913.-TYPE: MEX ico. : rocky soil, Cerro de Paxtle, vicinity of San Luis Tultitlanapa, near Oaxaca, Sep 1908, Purpus 3022 (holotype: GH!; isotypes: BM! F! MO! NY! UC! US!). Simsia sanguinea subsp. albida S. F. Blake, Contr. U.S. Natl. Herb. 22: 630. 1924.-TYPE: GUATEMALA.Huehuetenango: among stones on lightly wooded limestone hills, Uaxac Canal, Quen Santo, 1300 m, 21 Jul 1896, Seler & Seler 3098 (holotype: GH!; isotype:MO!). Aspilia grosseserrata M. E. Jones, Contr. W. Bot. 18: 84. 1933.-TYPE: MEX ico. Jalisco: La Barranca, Guadalajara, 17 Nov 1930 [16 or 23 Nov on some sheets], Jones 27778 (lectotype, here designated: POM!; isolectotypes: CAS! MO! NY! UC!). Cosmos hintonii Sherff, Brittonia 16: 72. 1964.-TYPE: MEXICO.Guerrero: Mina, Jiotes, grassy hill, 500m, 11 Sep 1937,Hinton et al. 10647 (holotype: MICH!; isotypes: G! GH! K! LL! UC! US!, photo: F!). 46 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 30

Herbaceous perennials to suffrutescent, taprooted perennials. Stems 0.3-1.5 m tall, 0.2-1 cm in diameter, or occasionally flowering at 0.1 cm in diameter, erect (rarely decumbent), green to brown or purple, usually harshly hispid and inter mixed glandular-puberulent. Leaves extremely variable, petioles subsessile to 6 cm long, 1-2 mm in diameter, winged throughout with wings to 3 cm wide, or only partly winged by the decurrent blade, or unwinged, often with stipular lobes at the base, these often dilated and cordate-clasping but not fused with those of adjacent leaves; blades 1.5-16 cm long, 0.5-18 cm wide, ovate to linear-ovate to linear, herbaceous, usually narrowly to deeply 3-lobed from the base, sometimes many lobed throughout into linear-pinnatifid segments, or sometimes unlobed, base cuneate and leading into the often winged petiole, apex acute to long-acuminate, usually rough scabrous andwith intermixed roughwhitish hispid vesture, the hispid vesture often confined to the veins, sometimes merely puberulent abaxially and scabrous adaxially, glands present or absent, if present usually denser abaxially, margins subentire tomore commonly dentate. Inflorescence corymbiform, loosely branched, often forming the upper one-half of the plant; peduncles 1 to more commonly 3-12 cm long, 0.5-1 mm in diameter, hirsute and glandular, sometimes puberulent. Heads radiate, 9-16 mm long, 15-25 mm in diameter. Involucre campanulate, 8-14 mm long, 6-10 mm in diameter; phyllaries 15-25, 2-3-seriate, unequal to nearly subequal, appressed to reflexed, acute to long-acuminate at the tips, usually green to greenish black, rarely tingedwith purple, strigose and hirsute, sometimes intermixed glandular-puberulent, margins entire; outer phyllaries 3.8 11mm long, 1-2.2 mm wide, lance-ovate; inner phyllaries 6.2-13 mm long, 1.2-2.5 mm wide, lanceolate to linear. Pales stramineous, 6.6-10.5 mm long, 1.2-2.5 mm wide, acute to acuminate, antrorsely pubescent apically. Ray florets 6-15; ligules 7.3-13.5 mm long, 3.7-5.5 mm wide, light pink to deep purple, rarelywhite or lemon-yellow; tube 1.2-2.3 mm long, 0.4-0.6 mm in diameter; ovary 4-6.7 mm long, 0.5-1 mm wide. Disc florets 17-44, corollas white (lemon-yellow when the rays are lemon-yellow), thewhite corollas turningpurple with age; corolla 5.5-7.5 mm long, 1-1.5 mm in diameter; tube 0.7-1.6 mm long, 0.5-0.8 mm in diameter; lobes 0.7-1.5 mm long, 0.5-0.9 mm wide; anthers 3.2-4.5 mm long, yellow, some times purplish on the distal ?4-?2, sometimes black immediately below the apical appendages, terminal appendages nearly black throughout or stramineous and black along the margins; achenes 3.2-6.5 mm long, 1.9-3.7 mm wide, laterally flattened, awns absent or 2.3-4.3 mm long, squamellae rarely present. Chromo some number: n = 17. Phenology. Flowering fromAugust throughFebruary. Distribution (Fig. 18). InMexico from the extensive barranca system about Guadalajara, Jalisco, into western Michoacain, throughout SW Mexico, , Guerrero, and Oaxaca to central Veracruz, and from Chiapas to Guatemala and El Salvador; open meadows and in shallow rocky soil in rocky areas or on cliffs, and along roadsides, in areas of tropical deciduous forests to oak-pine forests; 450 2480 m.

REPRESENTATIVE SPECIMENS. Mexico. CHIAPAS: 3 km S of Comitan near old airport, Breedlove & Davidse 54889 (CAS, MICH, TEX); 4 km above Motozintla on new rd to El Porvenir, Breedlove & Strother 46316 (DS); 36 mi W of Villa Flores, Cronquist & Sousa 10461 (DS, DUKE, ENCB, GH, MEXU, MICH, MSC, NY, TEX, US); Hacienda Monserrate, Purpus 9122 (GH, MO, NY); along Rt 1990 SIMSIA 47

1020 960 goo

20~

i S. sanguinea

960 90o

FIG. 18. Distribution of Simsia sanguinea.

190, 4.6 mi S of rd directly into town center of La Trinitaria, Spooner 2772 (MEXU, OS, UC). GUERRERO: Cutzamala, Hinton 6706 (F, GH, K, NY); Placeres-Cigarillo, Hinton 9818 (K, US); Rt 95, just S of Chilpancingo de los Bravos, Spooner & Burgos 2604 (COL, MICH, OS, TENN, UC, US, WIS); near dam, along rd from Chilpancingo de los Bravos to Ayotzinapa, Spooner & Burgos 2607 (MEXU, OS, TEX, UC, US).-JALISCO: La Barranca near Guadalajara, Pringle 1738 (BM, F, G, GH, LE, M, MASS, NY, S, TEX, UC, US, W), Pringle 4044 (M, MEXU), 11513 (CAS, F, GH, L, MEXU, MO, MSC, SMU, US); near bridge crossing of toll road over Rio Grande de Santiago, SE of Guadalajara, Spooner & Villasefior 2495 (GH, LE, MEXU, MICH, NY, OS, TEX, UC, WIS), Vil lasefior & Spooner 684 (MEXU, OS); Rt 41, 4 mi N of city limits of Guadalajara, Spooner & Villasefior 2505 (COL, CHAPA, F, GH, MEXU, MICH, NY, OS, OSH, P, TEX, UC, US, WIS); 8 mi SE of Tizapan el Alto near border with Michoacan, Spooner & Villasenior 2537 (BM, MICH, OS, OSH, P, UC, US, WIS), Villasefior & Spooner 723 (CAS, MEXU).-MExico: just NE of Temascaltepec along Rt 140, Anderson & Laskowski 3937 (DUKE, ENCB, F, MICH), Spooner & Villasenior 2572 (MEXU, MO, OS, P, TEX, US, WIS); Temascaltepac, Hinton 6706 (K); El Durazno, 4 mi SE of dam of lake by Valle de Bravo on rd to Colorines, Spooner & Villaseuior 2567 (G, MEXU, MICH, OS, US, WIS), Villasefior & Spooner 767 (CAS, MEXU); 1mi NE of rd from intersection of Valle de Bravo Rd & rd to , Spooner & Villasefior 2570 (MEXU, MO, NY, OS).-MICHOACAN: Tiquicheo, Hinton 13320 (ARIZ, ENCB, GH, K, LL, MICH, MO, TEX, US, W); Salto de Enandio, Hinton 13487 (ARIZ, LL, MICH, US); 12 km N of Huetamo, rd to Zitacuaro, Nuinez 6660 (TEX).-MORELOS: Xochicalco, Seler & Seler 5398 (GH); Jojutla, Miranda 1572 (MEXU).-OAXACA: 24 km NE of Sola de Vega along rd to Oaxaca, Breedlove 12254 (CAS, ENCB, MICH, NY, US); Xochimilco, Conzatti 3655 (MEXU); Cerro San Felipe, Conzatti 4045 (MEXU), Stuessy & Gardner 4271 (ASU, ENCB, OS); 10.1 mi NW of Tamazulapan on Rt 190, Crawford et al. 1305 (MEXU, MO, NY, OS, OSH, TEX, UC, US); 54 mi SE of Oaxaca at Portillo San Dionisio, Davidse & Davidse 9610 (MO); 12 mi S of Huajuapan de Le6n on Pan-American Hwy, Iltis et al. 1189 (TEX, WIS); S of Huajuapan de Le6n, SE of jct with Rt 125 to Tehuacan, Keil & Luckow 15491 (OBI, OS), Spooner & Dorado 2641a (MEXU, OS, UC, US, WIS); 7 mi N of Oaxaca, Kenoyer 1643 (GH); 16-17 mi S of Hwys 125 & 190, Lane & Olsen 2095 (TEX); Las Sedas, Pringle 5756 (GH); 3 km SE of Ocatlan, Solano & Vara 115 (CHAPA); S of Huajuapan de Le6n, SE of jct with Rt 125 to Tehuacan, Spooner & Dorado 2649b (MEXU, OS, TEX, UC, WIS); along Rt 190, 0.1 mi S of turnoff to Etla, Spooner & Villasenior 2658b (GH, OS, TEX, UC, US); Rt 195, 9 mi SE of Matatlan, 0.9 mi SE of intersection of rd to Ocotepec, Spooner 2819 (OS); 9 mi N of Pochutla, Turner 80A-25 (CAS, TEX).-PUEBLA: Km 17, on Tehuacan-Huajuapan de Le6n Rd, 48 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 30

23.5 mi N of Hwy 190, Bartholomew et al. 3043 (CAS); Km 84.5 on Tehuacdin-Huajuapande Ldon Rd, Panero & Schilling 651 (TENN); Caltepec, El Tecomite, El Gavilin, NW of San Sim6n, Tenorio & Romero 7558 (TEX).-VERACRUZ:Mirador, Consaquitla, Liebmann 492 (C, F, G, GH, L, LE, S, US), Sartorius 1146 (GH, P); Totutla, Encinal, Ventura 4121 (ASU, CAS, ENCB, MICH, TEX). Guate mala. BAJAVERAPAZ: above San Miguel Chicaj, Molina & Molina 27799 (ENCB, F, MICH, UC). CHIMALTENANGO:Almeda, Johnston 362 (F).-EscuINTLA: Aguilar 1748 (F).-GUATEMALA:large arroyo S of Guatemala on rd toAmatitlin, Standley 62834 (F, GH).-HUEHUETENANGO:along Rt CA 1, 2.3 mi SE of border with Mexico at La Mesilla, Spooner 2767 (F,MEXU, OS, US); Rt CA-1, ca. 35 mi WNW of Huehuetenango, Stuessy & Gardner 4316 (ASU, COL, ENCB, MO, OS, TEX, WIS). JALAPA:Montania Durazno, 2mi E of San Pedro Pinula, Steyermark32982 (F, US).-JUTIAPA: vicinity of Jutiapa, Standley 75023 (F).-SACATEPAQUEZ:6 mi N of Antigua, 0.1 mi N of intersection of rd to San Andres Itzapa, Spooner & Dorado 2684b (BM, F, LE, MEXU, OS, OSH, UC, WIS); 1.6 mi E of El Tejar on rd toGuatemala, Spooner 2761 (MEXU, OS).-SOLOLA: 3 mi NW of Panajachel, Spooner & Dorado 2681 (BM, F, LE, MEXU, MICH, OS, OSH, UC, US, WIS). El Salvador. SANTAANA: Chalchuapa, Calder6n 998 (US).

Simsia sanguinea exhibits the greatest range of morphological diversity of any species in the genus, but usually is readily distinguished by its pink to purple ligules and generally rough scabrous pubescence. It could with some justification be treated as three or four intergrading varieties. The common form of the plant is upright, 1-1.5 m tall, with harshly pubescent leaves that are deeply to narrowly 3 lobed from the base. On grassymeadows adjacent to and on rock ledgeswithin the extensive barranca system about Guadalajara (all plants listed from Jalisco), there occur populations distinguished by having leaves and heads at the large end of the size range for the species.Within this area, the plants show relatively little variabil ity, but similar forms are found elsewhere throughout the range of the species (e.g., Chiapas: Breedlove & Strother 46316, Conzatti & Gonzalez 899. Guatemala: Spooner 2767). In the Sierra Madre of northwestern Guerrero (Hinton 9818, Hinton et al., 10647, [the type of Cosmos hintonii Sherff]), immediately adjacent states ofMexico (Hinton 6706) andMichoacan (Hinton 13320, Nuiiez 660), from 450-500 m, there are populations distinguished by relatively thin, linear-pinnatifid leaves. Although somewhatmorphologically distinct and geographically isolated,most of these collec tions are from isolated areas and little is known about populational variability or intergradation of these features. Occurring throughout parts of Oaxaca (e.g., Conzatti 3655, 4045; Davidse & Davidse 9610; Kenoyer 1643; Pringle 9756; Solano & Vara 115; Spooner 2819; Stuessy & Gardner 4271) from 1500-1800 m is a usually low-growing form, 0.5 m or less tall, sometimes with decumbent stems, and with small, narrow, sometimes linear leaves with a conspicuous harsh whitish vesture. The extensive sympatric intergradationwith other populations, however, suggests that these are best re garded as only extreme variants worthy of mention but not formal recognition. In southern Puebla and immediately adjacent northwestern Oaxaca, there are populations distinguished solely by lemon-yellow corollas of both ray and disc florets, referred to below as "yellow-flowered" (e.g., Oaxaca: Crawford et al. 1305, Iltis et al. 1109, Keil & Luckow 15491, Lane & Olsen 2098, Spooner & Dorado 2641a, 2649b, Webster 16984. Puebla: Bartholomew et al. 3043, Panero & Schilling 651, Purpus 3022 [the type of S. triloba], Tenorio & Romero 7558). Within this area, these variants apparently completely replace the more common populations that possess light pink to purple ligules and white disc florets, referred to below as 1990 SIMSIA 49

"pink-flowered". These yellow-flowered variants frequently are misidentified as S. amplexicaulis, which differs by its annual habit, orange-yellow ligules, and leaves not as rough-scabrous as those of S. sanguinea. Plants with white ligules and white disc florets occasionally are found intermixed both in these yellow-flowered popula tions (e.g., Puebla: Bartholomew et al. 3045), and in the pink-flowered populations (e.g., Mexico, Guerrero: Spooner & Burgos 2607. Chiapas: Breedlove & Davidse 54889, Purpus 9122. Guatemala. Huehuetenango: Seler & Seler 3098 [the type of S. sanguinea subsp. albida]. Solola: Spooner & Dorado 2681). There ismuch vegeta tive diversity within the white- and yellow-flowered populations of S. sanguinea, similar to that found within the pink-flowered ones. For example, Spooner & Dorado 2649b is a typical, upright, broad-leaved plant, but Iltis et al. 1189 and Spooner & Dorado 2641a are prostrate, linear-leaved plants similar to others in Oaxaca and other areas (but with pink ligules). Crosses between the pink, yellow, and white color forms (see Appendix 2) produced 34 flowering F1 individuals. The corollas of the disc and ligule flowers of these hybrids varied in color from the extreme parental conditions to various combi nations of colors restricted to each type of corolla, i.e., the disc corollas were white, yellow, or intermediate, but not pink. Sometimes the ligules were streaked with both pink and yellow. My initial intent was to segregate the yellow-flowered plants of S. sanguinea because of the apparent replacement of the pink-flowered populations by the yellow-flowered ones in southern Puebla and adjacent northwestern Oaxaca. I have decided against recognizing the yellow-flowered plants at the varietal level, how ever, because flower colors are the only characters that distinguish these popula tions, and because white-flowered plants occur within both the yellow- and pink flowered populations. Also, the great vegetative diversity foundwithin the range of the yellow-flowered populations parallels that found within much of the pink flowered populations. The protologue of Aspilia grosseserrata lists 17 Nov 1930 for the date of the type collection, but Jones's specimens bearing identical data differ by the dates: 16 Nov, POM; 17 Nov, MO, UC; and 23 Nov, CAS, NY. According to Lenz (1986), Jones's field notes indicate he actually collected the type on 15 November. Lenz also documents, however, that Jones visited La Barranca on all of the dates listed above as well. Whether these sheets represent separate collections or one collection later confused by labelling errors is unclear. They all appear similar and apparently represent the same collection, and here all are regarded as type material.

12. Simsia calva (A. Gray & Engelmann ex A. Gray) A. Gray, Boston J. Nat. Hist. 6 (P1. lindheim. 2): 228. 1850. Barrattia calva A. Gray & Engelmann ex A. Gray, Proc. Amer. Acad. Arts 1: 48. (Dec 1846 or Jan 1847 fide Stafleu & Cowan, 1976). Encelia calva (A. Gray & Engelmann ex A. Gray) A. Gray, Proc. Amer. Acad. Arts 8: 658. 1873.-TYPE: U.S.A. Texas: rocky soil, in open woods between the headwaters of the Guadalupe & Pedernales Riv ers, Oct 1845, Lindheimer 432 (lectotype, designated by Blake, 1913:GH!; isolectotype:MO!). Simsia subaristata A. Gray, Mem. Amer. Acad. Arts 1 (P1. fendl.): 84. 1849. Encelia subaristata (A. Gray) A. Gray ex Hemsley, Biol. cent.-amer., Bot. 2: 185. 1881. Simsia calva var. subaristata (A. Gray) S. F. Blake, Proc. 50 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 30

Amer. Acad. Arts 49: 379. 1913.-TYPE: MEXICO.Nuevo Leon: Bishop's Hill near Monterrey, 6 Feb 1847, Gregg 47 (lectotype, designated by Blake, 1913:GH!; isolectotypes: MO[2]! NY!).

Herbaceous perennials to suffrutescent perennials arising from a somewhat fleshy, fusiform-thickened root up to 11 cm long and 3.5 mm in diameter. Stems 0.3-1.5 m tall, 0.2-1 cm in diameter, ascending to erect, green to purple, scabrous, often hirsute and glandular-puberulent. Leaves with petioles 0.2-3 cm long, 0.5-1 mm in diameter, at base with stipules of opposite leaves fused to form nodal discs 0-1.5 mm long and wide; blades 2-8 cm long, 1.5-6 cm wide, ovate, herbaceous, unlobed to deeply 3-lobed, base cordate to truncate or cuneate, apex acute to acuminate, eglandular, scabrous and hirsute, margins crenate-dentate, rarely subentire. Inflorescence usually of solitary heads (sometimes 2-3 in corymbs) on peduncles 3-30 cm long, arising from many nodes, 1-2 mm in diameter, hirsute, sometimes scabrous and glandular-puberulent. Heads radiate, 11-16 mm long, 20 40 mm in diameter. Involucre campanulate, 10-12 mm long, 7-16 mm in diameter; phyllaries 21-43, 3-seriate, unequal to subequal, appressed tomoderately to greatly reflexed, acute to attenuate at the tips, green to yellow-green or green-black, rarely sinuate, scabrous and hirsute, sometimes glandular,margins entire; outer phyllaries 4.5-12.3 mm long, 1.5-3 mm wide, lance-ovate to lanceolate; inner phyllaries 6.4 13mm long, 1-2.5 mm wide, lanceolate to linear or oblanceolate. Pales 5.5-10 mm long, 1.5-2.5 mm wide, stramineous, sometimes purple apically, acute to acumi nate, scabrous apically. Ray florets 8-21; ligules 5-16 mm long, 2-6.5 mm wide, light orange-yellow adaxially, often lined or sometimes entirely brown or purple abaxially; tube 1-2.1 mm long, 0.3-0.6 mm in diameter; ovary 3-5 mm long, 0.5 1.1 mm wide. Disc florets 26-154, usually more than 90; corolla 5-7 mm long, 1 1.5 mm in diameter; tube 0.5-1.2 mm long, 0.4-0.8 mm in diameter; lobes 0.8-1.3 mm long, 0.6-0.9 mm wide; anthers 3.1-4.3 mm long, yellow, rarely black, termi nal appendages stramineous; achenes 3.5-5.7 mm long, 1.8-4.6 mm wide, laterally flattened, pappus commonly absent or of awnsminute to 4 mm long. Chromosome number: n = 17. Phenology. Flowering throughout the year. Distribution (Fig. 17). In theU.S.A. widespread throughout central, southern, and southwestern Texas from the south Texas Plains to the trans-Pecosmountains, SE , and in Mexico mostly on the lower to upper slopes of the Sierra Madre Oriental to the northern part of the Altiplano; in sand to heavy clay soils, or in rock crevices, often in limestone, on open prairies or thickets or oak savannas, or in rocks along streams, often along roadsides, to upland pine or pine-oak forests; 30-2400 m.

REPRESENTATIVESPECIMENS. U.S.A. NEWMEXICO: Lincoln Co., Devil's River, El Capitan Mtns, 22 Jul 1900, Earle & Earle s.n. (NY).-TEXAS: ATASCOSACO., Pleasanton, Palmer 9790 (DS, MO, S, US). Baylor Co., Seymour, Clark 4695 (G). Bexar Co., rocky hills of the upperGuadalupe River, Lindheimer 433 (BM, CGE, G, GH, GOET, K, LE, MICH, MO, NY, OXF, SMU, UC, US, W); Comanche Spring, New Braunfels, etc. Lindheimer 900 (ARIZ, BM, C, F, G, GH, LE, M, MEXU, MO, NY, SMU, TEX, US, W). Blanco Co., 4.7 mi N of Blanco on US 281, Flyr 472 (DUKE, SMU). Brewster Co., W slopes of Baldy Peak, Glass Mtns, Warnock W 264 (ARIZ, DS, GH, LE, MO, NY, PH, S, SRSC, TEX, UC); Sul Ross Campus, Paul 1 (SRSC). Brown Co., Brownwood, Palmer 26793 (MO). Burnet Co., 3.1 mi N of Burnet, Cory 15736 (GH). Callahan Co., Rt. 80, 2 mi W of Baird, Spooner & 1990 SIMSIA 51

Schilling 2385 (BM, COL, ENCB, LE, MEXU, MICH, MO, OS, P, WIS). Cameron Co., 2 mi NE of Combes, Shinners 17747 (SMU, TEX). Coke Co., 5 mi N of Sanco, Shinners 8405 (SMU). Coleman Co., Santa Ana Mtns, Pruitt 233 (TEX). Concho Co., 5.6 mi E of Eden on US 87, Stuessy & Meacham 3511 (NY, OS). Coryell Co., N of Killeen, Wolff 3273 (SMU, US). Crockett Co., 8 mi W of Ozona on US 290, Urbatsch 965 (LL). Culberson Co., Hurd's Draw, Janszen 61 (TEX). Dimmit Co., 3mi S of San Diego on Hwy 1329, Mahler 5277 (SMU, WIS). Duval Co., 4 mi SE of Bruni, Alvarez et al. 7611 (DUKE, LL, TEX). Edwards Co., 30 mi NW of Rocksprings at substation 14, Cory 51726 (DS,MICH, NY, SMU, US). Frio Co., 6 mi N of Pearsall, Shinners 7430 (SMU, TEX, UC). Gillespie Co., Spring Creek, Jermy 863 (F). Glasscock Co., 10mi W of Garden City, Turner 4953 (TEX). Goliad Co., 0.5 mi W of Goliad, Cory 55138 (SMU, US). Hamilton Co., 0.3 mi SW of IndianGap, Cory 53791 (SMU). Hays Co., SanMarcos, Palmer 12119 (G, GH, MO, NY, TEX). Hidalgo Co., 3mi S ofWeslaco, Tidwell 111 (TEX). Howard Co., Big Spring State Park, Correll 16682 (SMU). Jeff Davis Co., JeffDavis Mtns, Rt 118, 1mi NW of rd into park, 4.8 mi from St Rt 17, Spooner & Schilling 2391 (BM, CHAPA, F, MEXU, MO, NY, OS, US, WIS). Jim Hogg Co., ca 36 mi S of Hebbronville on Rt 1017 near Starr Co. line, Strother 860 (NMC, UC). Jones Co., W shore of Fort Phantom Lake, 9 mi N of Abilene, Hender son 63-699 (SMU). Karnes Co., 7 mi NW of Karnes City, Turner 4583 (TEX). Kendall Co., 0.8 mi SW of Bergheim on Hwy 46, Shinners 32924 (SMU). Kerr Co., Kerrville, Heller 1860 (G, NY, UC, US). Kimble Co., Junction, Cory 49084 (CAS). Kinney Co., 4 mi NE of Brackettville, Strother 268 (DS, ENCB, SMU, TEX). Kleberg Co., Kingsville, Benke 5456 (F, G, GH). Lampasas Co., Mountain View School, 12 Jun 1941, Tharp s.n. (TEX). Live Oak Co., 8 mi S of George West on Rt 281, Flyr 350 (NY). Mason Co., Mason-Eden, 31 Aug 1929, Whitehouse s.n. (TEX). Maverick Co., 12 mi E of Eagle Pass, 20 May 1898, Bray s.n. (TEX). McCulloch Co., 1 mi S of Brady on rd to Mason, Gould 5691 (SMU, TEX). McMullen Co., W of Three Rivers on Henderson Ranch, Whitehouse 11027 (SMU). Medina Co., 3 mi W of Castroville, Spooner & Brandenburg 2177 (OS). Menard Co., 4.2 mi S of Menard on US 83, Raven & Gregory 19266 (DS, TEX). Mills Co., 3.2 mi SE of Goldthwaite, Shinners 16830 (SMU). Mitchell Co., NE ?4, Sect 17. S.P.R.R. Block 18, Pohl 4627 (S, SMU). Nolan Co., Sweetwater, Palmer 14539 (MO). Pecos Co., Tunis Creek at East Escondito Springs, Tharp 43-903 (MO, NY, PH, TEX, UC). Presidio Co., Vieja Pass, Vieja Mtns, Warnock 8887 (MICH, SRSC). Real Co., Barksdale, Connell 6506 (LL, TEX). Reeves Co., along canal at Balmorhea, Fletcher 600 (SRSC). Runnels Co., 10 mi NW of Ballinger, Shinners 31753 (TENN, SMU). San Patricio Co., Lake Corpus Christi State Park near Mathis, Traverse 1788 (F, GH, LL, SMU, TEX). San Saba Co., Chappel, Barnette 275 (SMU). Schleicher Co., Rt. 277, 8.5 mi N of Eldorado, Raven & Gregory 19248 (DS). Scurry Co., 5 mi N of Fluvanna, Correll & Johnson 17198 (LL). Shackelford Co., 15 mi W of Albany, Whitehouse 16717 (MICH, SMU). Starr Co., 6 mi N of Rio Grande City, Clover 1672 (LL, MICH). Sterling Co., 1.5 mi NE of Sterling City, Correll 31722 (LL). Sutton Co., 7.3 mi N of Sonora, Cory 50555 (MICH, SMU). Taylor Co., 13 mi S of Merkel of Hwy 126, Mahler 3007 (SMU). Terrell Co., Rt 90, 6.7 mi E of Sanderson, Raven & Gregory 19203 (DS). Tom Green Co., 9 mi N of San Angelo on Hwy 277, Watson 540 (ASU, TEX). Travis Co., Austin, Hall 339 (G, GH, MO, NY, US). Upton Co., 4.1 mi N of Rankin, Shinners 31920 (SMU). Uvalde Co., 21 mi N of Uvalde, Shinners 7315 (DS, SMU, WIS). Val Verde Co., View point, Amistad Recreation Area along Rt 90, ca 3 mi E of dam, Urbatsch 928 (LL). Webb Co., Laredo, Magnon & Rodriguez 31 (ENCB, TEX). Willacy Co., S of Raymondville, Gentry 710 (ARIZ, SMU). Zapata Co., 4 mi E of Zapata, Ariaza 55 (SMU, TEX). Zavala Co., 15 mi N of Crystal City, Shinners 7376 (SMU). Mexico. CHIHUAHUA:Chihuahua, Paray 2565 (ENCB).-:La Cuesta del Plumo on the Muzquiz-Boquillas Hwy, Chiang et al. 9211 (TEX); 1.1 mi N of rd to Laguna Churince along Cuatro Cienegas Hwy, Lehto et al. 5233 (ARIZ, ENCB, LL, NY); Saltillo, Palmer 615 (MASS, GH, US); 0.5 mi N of Toro Viejo on Rt 57, Reeves & Pinkava 13016 (ASU, OS); 14 mi N of Sabinas on Hwy 57, Spooner & Brandenburg 2181 (CHAPA, OS); 4 mi N of Nueva Rosita on rd to Melchor, Stuessy 909 (ENCB, NY, TEX); Rancho Santa Teresa, S of Castanios,Wynd & Mueller 177 (ARIZ, GH, K, NY, S, TEX, US).-NUEvo LE6N:Mamulique Pass, 70 km N ofMonterrey, Cronquist 11299 (MEXU, MICH, NY), Spooner & Triplehorn 1058 (CHAPA, LE, NY, OS, OSH, P, RSA, WIS); 2 km S of Puerto Calvertas, 46 km N of Dr. Arroyo, Flyr 1558 (TEX); Sierra Madre above Monterrey, Pringle 8739 (BM, C, CM, ENCB, F, G, GH, GOET, LE, LL, M, MEXU, MO, MSC, NY, S, UC, US, W); 18 mi W of Linares, Roberts & Keil 10456 (F, NY, OS); Linares-Iturbide Rd, 13 mi W of Rt 85, Spooner 2482 (ENCB, F, LE, MEXU, MICH, MO, OS, TENN, TEX, UC, US, WIS)-:11 mi N of Jimdnez, Johnston 4892 B (TEX); 17mi E of Victoria, Johnston 5785 (MEXU, MICH, TEX); 4 mi S of Jaumave, Stanford et al. 2287 (DS, GH, MICH, MO, NY, SMU, TEX, UC, US); 12 mi S of Laredo, Urbatsch 2407 (OS). 52 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 30

Simsia calva is easily identified by its fleshy, fusiform-thickened root, profuse basal branching, and solitary heads, and is generally not confused with any other species. The variability of awns and pubescence on the achenes of S. calva ledAsa Gray at various times (1846; 1849; 1850; 1873) to position this species under three genera and divide it into two species. As inmost other species of Simsia, these features often vary within single populations and are not recognized here. Two specimens of Gregg atMO bear identical data about the locality, date, and collector, but bear the number 147 instead of 47. I consider the number 147 a transcriptionalerror and these specimens as isolectotypes of S. subaristata.

13. Simsia lagascaeformis de Candolle, Prodr. 5: 577. 1836. Encelia lagascaeformis (de Candolle) A. Gray ex Hemsley, Biol. cent.-amer., Bot. 2:184. 1881. TYPE:MEXICO, 1883, Mairet s.n. (holotype: G-DC, microfiche: IDC 800.946.11.8!, photos: GH! MICH! MO!). Simsia exaristataA. Gray, Smithsonian Contr. Knowl. 5, Art. 6 (PI.wright. 2): 87. 1853. Encelia exaristata (A. Gray) A. Gray ex Hemsley, Biol. cent. amer., Bot. 2: 183. 1881. Simsia exaristata var. epapposa S. F. Blake, Contr. Gray Herb. 52: 45. 1917, nom. superfl.-TYPE: MEXICO.Sonora, or U.S.A. :Cochise Co.: valley of a tributaryof the San Pedro, Sep 1851,Wright 1224 (lectotype, here designated: GH!). Encelia purpurea Rose, Contr. U.S. Natl. Herb. 1: 336. 1895.-TYPE: MEXICO. Colima: Colima, in a creek bottom, 9 Jan-6 Feb 1891, Palmer 1105 (holotype: US-208967!; isotypes: BM! GH! K! LL! NY! US!). Encelia pilosa Greenman, Proc. Amer. Acad. Arts 39: 111. 1903.-TYPE: MEX iCO.Oaxaca; 17Oct 1899,Holway 3747 (lectotype, designated byMcVaugh, 1984:GH!). Simsia exaristata var. perplexa S. F. Blake, Contr. Gray Herb. 52: 45. 1917. TYPE: U.S.A. Texas: [El Paso Co.], valleys in the mountains E of El Paso, 12-17 Sep 1849,Wright 331 p.p. (holotype: K, tracingand fragment:GH!).

Taprooted annuals, but occasionally suffrutescent at base. Stems 0.2-4 m tall, 0.2-1.0 cm in diameter (or rarely up to 8 cm in diameter in persisting suffrutescent individuals), erect, sometimes decumbent at the base and rooting at the nodes, green to yellow, brown, or purple, glandular-puberulent, sometimes intermixed pilose or strigose. Leaves with petioles 1-7 cm long, often long-pilose at base, 1-2 mm in diameter, without wings, or very rarely with petioles ampliate at the base; blades 2-21 cm long, 1-16 cm wide, ovate to deltate, herbaceous, usually unlobed, rarely narrowly to deeply 3-lobed, base cordate to truncate or cuneate, apex acute or acuminate, indument variable, often scabrous and thinly hirsute, or sometimes puberulent to pilose, usually eglandular but sometimes with stipitate glands, mar gins crenate to dentate. Inflorescence corymbiform, tightly to loosely branched; peduncles 0.5-10 cm long, 1-1.5 mm in diameter, glandular-puberulent and hir sute, sometimes eglandular. Heads radiate, 10-14 mm long, 13-25 mm in diameter. Involucre campanulate, 8-12 mm long, 5-10 mm in diameter; phyllaries 13-19, 2 3-seriate, unequal, appressed or slightly reflexed apically, acute to acuminate at the tips, light green to yellow with purple margins or totally purple, usually glandular puberulent and pilose, sometimes eglandular, margins entire; outer phyllaries 2.2 1990 SIMSIA 53

960 9o0

0~~~~0

240~~~~ 24~~~~~~~~~~~~~~~~~~~~~~4

A S. eurylepis _ ,, * S. lagascaeformis

FIG. 19. Distribution of Simsia chaseae, S. eurylepis, and S. lagascaeformis.

6.7 mm long, 1.5-2.5 mm wide, ovate to lance-ovate; inner phyllaries 4.6-9.5 mm long, 1.5-2.7 mm wide, lanceolate. Pales 5.5-9.4 mm long, 1.4-2.5 mm wide, stramineous and purple apically or purple throughout, often nearly truncate api cally with an apical mucro, to acute to acuminate, antrorsely pubescent apically. Ray florets 5-10; ligules 5.1-12 mm long, 2.3-5.3 mm wide, orange-yellow; tube 1 1.7 mm long, 0.4-0.5 mm in diameter, ovary 3.3-6 mm long, 0.3-0.8 mm wide. Disc florets 13-27; corolla 5.8-6.7 mm long, 0.8-1.2 mm in diameter; tube 0.8-1.2 mm long, 0.5-0.8 mm in diameter; lobes 0.8-1.2 mm long, 0.5-0.8 mm wide; anthers 3.2-4 mm long, yellow proximally, usually purple (often drying bronze) distally, with terminal appendages stramineous; achenes 4.2-6 mm long, 2-3.5 mm wide, laterally flattened, awns 2.5-4.6 mm long or rarely absent, squamellae ab sent. Chromosome number: n = 17. Phenology. Flowering specimens can be found throughout the year, but most plants flower from September throughDecember. Distribution (Fig. 19). In theU.S.A, in SArizona, S New Mexico, SW Texas, inMexico south to central Oaxaca, also in Guatemala; mostly in areas of deserts or 54 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 30

tropical deciduous forests, along roadsides, in open fields, and along borders of agricultural fields; 140-2200 m.

REPRESENTATIVESPECIMENS. U.S.A. ARIZONA:Cochise Co., Tombstone, Peebles et al. 3377 (ARIZ, LL), Mason A-227 (CHAPA).-NEW MEXICO: Hildalgo Co., 4 mi SE of jct of NM Rt 81 along dirt rd in Hachita Valley, Keil et al. 13442 (OBI).-TEXAS: W Texas, valley of the Rio Grande, below Donada, Parry et al. 561 (NY, US). Brewster Co., Alpine, Steiger 938 (NY). Mexico. CHIHUAHUA:valley near Chihuahua, Pringle 321 (BM, DS, G, GH, K, MASS, MICH, NY); N of Jimenez, 5.1 mi N of jct of Rts 45 & 49, on rd to city of Chihuahua, Spooner & Brandenburg 2336 (CHAPA, OS, UC, WIS); Hwy 16, 17 mi NE of town square of Aldama, Spooner & Schilling 2456 (BM, COL, G, GH, LE, MEXU, MICH, NY, OS, OSH, TEX, UC, US, WIS); Rt 49, 9 mi S of S limits of Cd. Camargo, Spooner & Schilling 2458 (MEXU, OS, UC, WIS).-CoAHuILA: 32 mi NE of San Pedro, on Hwy 30, Henrickson 6009a (LL); Llano de Guaje, on rd from Santa Elena mines NE to Tanque La India, Johnson & Mueller 791 (MEXU, MICH, GH, LL).-CoLIMA: Colima, Palmer 1105 (BM, K, GH, LL, NY, US).-: Rt 49, 1mi NW of Peronal, Spooner & Schilling 2461 (LE,MEXU, OS, TEX, WIS).-GUANAJUATO:Mpio. Yuriria, Ejido Parangarico, Garcia Z-723 (CAS).-GUERRERO:Coyuca, Hinton 5549 (G, GH, K, MO, NY); 15mi N of Zumpango del Rio on Rt 95, 8.4 mi S of Rio Mezcala, Keil & Luckow 15331 (OBI, OS); Chilpancingo de los Bravos-Chichihualco Rd, 6 mi W of Rt 95, Spooner & Burgos 2603 (BM, LE, MICH, OS, OSH); Rt 95, on S side of Chilpancingo de los Bravos, Spooner & Burgos 2606 (F, MEXU, MO, NY, OS, TENN, TEX, UC, W); 10-30 mi S of Sabana Grande on Rt 95 on rd to Chilpancingo, Stuessy & Gardner 4211 (ASU, ENCB, MO, OS, WIS).-HIDALGO: Jacala, Kenoyer 594 (MO).-JALISCO: Rt 80, 6 mi SW of San Juan de los Lagos, Spooner & Villasefior 2489 (C, GUADA, LE, MEXU, MICH, NY, OS, TENN, TEX, RSA, UC, US).-MIxico: Pungarancho, Hinton 8578 (GH, LL, MICH).-MICHOACAN: San Gregorio, Villarreal 196 (ENCB).-MORELOS: S side of Cuernavaca on free rd to Acapulco, Spooner & Burgos 2600 (MEXU, OS, TEX, US, WIS); S of Cuautla, 1.7 mi N of rd to Emiliano Zapata, Spooner2836b (MEXU, OS, US); 10-20 mi S of Cuernavaca on toll rd 95-D, Stuessy & Gardner 4239 (ASU, ENCB, MO, OS, WIS).-OAXACA: Rt 190, 14 mi S of Nochixtlan, King 2518 (DS, MICH, US); jct of Rts 131 and 190 in Cd. Oaxaca, Spooner & Triplehorn 2098 (CHAPA, OS, TEX, US, WIS); Rt 190, 7.1 mi S of rd to Chicachua, S of Ascunci6n Nochixtlan, Spooner & Dorado 2656 (MEXU, OS, US, WIS); along dirt rd from Jayacatlan to La Uni6n, 8 mi N of Jayacatlan, Spooner2831 (MEXU, OS); 1mi N of Monte Alban Park on rd to Cd. Oaxaca, Stuessy & Gardner 4262 (OS, TEX, WIS).-PUEBLA: 20 mi N of Huahuapan de Le6n, Cronquist & Sousa 10403 (ENCB, MEXU, MICH, NY); Cuautla-Matamoros Rd, 9 mi E of Tepexco, Spooner & Dorado 2627 (MEXU, OS, WIS); 4 mi N of Tehuitzingo, Spooner & Dorado 2636 (OS, OSH, UC, US).-SAN LuIs POTOSI:Rfo Verde, Calvert 123a (TEX); El Quelital, Rzedowski 6828 (ENCB, MEXU, MICH, WIS).-VERACRUZ: Orizaba, Botteri 804 (GH), Bourgeau 3320 (G, K), Mueller 143 (NY), 240 (NY, US), 597 (NY).-SONORA: Sonora, Thurber 953 (GH, UC). Guatemala. CH1 MALTENANGO:Almeda, Johnston 987 (F).-GUATEMALA.Antigua, Heiser R5 (IND); caldera, Volcdn de Pacaya, Tonduz 508 (MO).-SACATEPtQUEZ:near Antigua, Standley 58037 (F,NY), 60374 (CAS, F, U). Putative hybrid between S. lagascaeformis and S. amplexicaulis: JALISCO: Rt. 80, 6 mi SW of San Juan de Los Lagos, Spooner & Villasefior 2491 (MEXU, OS).

Simsia lagascaeformis usually is readily distinguished by its usually annual habit, unwinged petioles often long-pilose at the base, unequal phyllaries, orange ligules, and anthers yellow proximally andwith a purple band (often drying bronze) distally. It bears much overall resemblance to S. eurylepis, which is eradiate and lacks the purple distal band of the anthers. It also bears some resemblance to S. holwayi, which is perennial, has nodal discs, and lacks the purple distal band on the anthers. The nomenclatural history of S. lagascaeformis is complex, largely because of the great emphasis that Gray and Blake placed on the pubescence and presence/ absence of awns of the disc achenes. This species, as many others in Simsia, exhibits polymorphism for these traitswithin single populations (seeMorphology). Simsia lagascaeformis was described with glabrous achenes. Gray (1853) later described S. exaristata, based on Wright 331 p.p. (that part of the collection with calvous 1990 SIMSIA 55 achenes), and Wright 1224. Blake (1913) learned (by a letter of de Candolle) that the type collection of S. lagascaeformishas pubescent, awned achenes, not glabrous as originally described, and used these achenial characters to distinguish the two species. Blake (1917) later noted the polymorphic nature of this trait inWright 331, and described S. exaristata var. perplexa, based on that portion of this collection with pubescent, awned achenes. Blake (1917, p. 45) realized the tenuous nature of the features separating S. lagascaeformis and S. exaristata, but maintained them as weak segregates on the basis of other characters, the "longer narrower phyllaries, narrower pales, longer disc florets, fewer larger heads, and leaves not at all glandu lar beneath" of S. exaristata. A critical examination of these features indicates that they vary considerably throughout the range of the species and are not taxonomi cally useful. The only discernible trend is for some plants in the northern part of the range to have less gradate phyllaries. Although this is pronounced in some individu als, it is far from constant and specimens with less gradate phyllaries are also found farther south. In addition to this nomenclatural confusion, there are problems regarding the localities for the types of these names. Wright's collections have partial or ambigu ous locality-data, which are complicated by subsequent place name or boundary changes. Another ambiguity is that even thoughWright maintained his own collec tion numbers, these rarely appear on his collections, because the specimens were subsequently distributed by Gray under new numbers. Also, these new distribution numbers may incorporate more than one of Wright's collections. Shaw (1987) provided data that help clarify the collection localities of Wright 331 and 1224. Because no more than one locality is mentioned in each of the respective protologues, it is very possible that each number represents a single collection. All collections made between 12-17 Sep were from the mountains of El Paso, El Paso Co., Texas. The locality of Wright 1224 is ambiguous, as the San Pedro River traverses the present United States/Mexican boundary andWright's exact itinerary during these dates is unknown. Because of the confusion concerning Wright's collections, and the variation in achene pubescence, only holotypes or lectotypes are here designated for their respective names, and the others are considered possible isotypes, but not so anno tated. They are: Wright 331: BM, GH, OXF; Wright 1224: CGE, G, GH, MO, NY, PH, US, photo: TEX. Simsia lagascaeformis is almost always an herbaceous species. In some habitats, however, such as along irrigation ditches (e.g., Spooner 2831, 2836b), the plants may persist and the base may increase in diameter and become woody. Spooner 2836b had a woody base to 8 cm in diameter. The leaves of this species are usually unlobed, without wings on the petioles, and lack nodal discs. Rarely, lobed leaves and/or petioles with basal wings or nodal discs occur (e.g., some duplicates of Spooner & Dorado 2646). Simsia lagascaeformis apparently hybridizes naturally with S. amplexicaulis. Intermediates (Spooner & Villasefior 2491) occur with S. lagascaeformis (2489) and S. amplexicaulis (2490). The intermediates possess the amplexicaul leaves of S. amplexicaulis and the purple, subequal phyllaries of S. lagascaeformis. Rose (1895) designated Palmer 1105 as the type of Encelia purpurea. Blake (1913, p. 383) commented that this species "seems to be merely a peculiarly branched and perhaps somewhat teratological condition, with very numerous 56 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 30

capitula and somewhat flattened branches." The branches are mostly terete, and this collection has a normal morphology for S. lagascaeformis. Its small leaves suggest it is new growth from a damaged plant or perhaps a dwarf individual.

14. Simsia eurylepis S. F. Blake, Proc. Amer. Acad. Arts 49: 382. 1913.-TYPE: MEXICO.San Luis Potosi: Distr. Ciudad delMaiz, near Gallinas, Feb 1888, Seler & Seler 684 (holotype: GH!; isotypes:MO! US!). Simsia submollicoma S. F. Blake, Proc. Amer. Acad. Arts 49: 381. 1913. TYPE:MEXICO. Tamaulipas: vicinity of Tampico, 10Mar-18 Apr 1910, Palmer 250 (holotype: GH!; isotypes: BM! F! K! MO! NY! US!).

Taprooted annuals. Stems 0.5-3 m tall, 0.5-1 cm in diameter, erect, green to purple, glabrous to sparsely pilose. Leaves with petioles 1-8 cm long, 1 mm in diameter, at base often long-pilose, unwinged or with stipules minute to 3.2 cm long, these often united from opposite leaves to form nodal discs; blades 3-15 cm long, 2-13 cm wide, broadly to narrowly ovate, herbaceous, unlobed to deeply 3 lobed, base cordate to truncate or cuneate, apex acute to acuminate, puberulent, sometimes intermixed with hispid vesture and longer pubescent vesture, margins dentate to crenate. Inflorescence corymbiform to paniculate, tightly to loosely branched; heads subsessile or borne on peduncles up to 10 cm long, 1 mm in diameter, sparsely tomoderately pilose and puberulent. Heads eradiate, 13-16 mm long; involucre urceolate, 10-13 mm long, 6-12 mm in diameter; phyllaries 19-32, 3-seriate, unequal, appressed, purple and green, moderately to densely sericeous to pilose, margins entire; outer phyllaries 3.7-6.5 mm long, 1.6-2.5 mm wide, ovate; inner phyllaries 8.7-10.7 mm long, 1.5-2.6 mm wide, ovate to lanceolate. Pales 7.4-9.8 mm long, 1.8-3.1 mm wide, stramineous to purple, acute to acuminate, antrorsely pubescent apically. Disc florets 19-41; corollas 6.3-8.5 mm long, 1.1 1.3 mm in diameter, orange-yellow; tube 1.5-2.5 mm long, 0.6-0.7 mm in diame ter; lobes 1.5-1.8 mm long, 0.6-0.7 mm wide; anthers 2.6-3.6 mm long, yellow, terminal appendages stramineous; achenes 4.2-5.8 mm long, 2.2-3.6 mm wide, laterally flattened, awns absent or 0.3-5 mm long, squamellae absent. Chromo some number: n = 17. Phenology. Flowering throughout the year. Distribution (Fig. 19). Along the Gulf coastal lowlands from Tamaulipas to ; common along roadsides and in old fields; 0-825 m.

REPRESENTATIVE SPECIMENS. Mexico. CAMPECHE: along Campeche-Uxmal Rd, 0.5 mi E of jct with Rt 180, Spooner 2799* (LE,MEXU, OS, P, TEX, WIS); 3mi W of Tikinmul, Spooner 2800* (MEXU, MO, OS, WIS); Ichek, Spooner 2801* (MEXU, OS, TEX); W side of Escarcega, Spooner 2812* (MEXU, NY, OS).-NUEvo LE6N: SE side of El Cercado, Spooner 2481* (CHAPA, COL, ENCB, F, LE, MASS, MEXU, OS, RSA, TENN, TEX, UC, US, WIS).-SAN LuIs POTOSf:valley of Rio Tampa6n, Chase 7494* (ARIZ, CAS, F, GH, MICH, MO, NY); 8 mi S of Valles, Crutchfield & Johnston 5206* (MEXU, MICH, TEX); Cd. Valles, Hansen & Nee 7359 (MO); 9 mi E of Cd. Valles, McGregor 16325 (MSC); El Abra, Rzedowski 7440 (ENCB, MEXU, MICH); N of Cd. Valles on Rt 85, 5.1 mi N of jct of Rt 86, Spooner & Brandenburg 2209* (GH, MEXU, OS, TEX, US, WIS); 30-40 mi W of C. Valles on Rt 86, Stuessy & Gardner 4039* (ASU, CHAPA, ENCB, MO, OS, WIS); 25mi E of Rio Verde, Stuessy & Gardner 4041 * (ASU, ENCB, MO, OS, WIS); 12mi E of Cd. Valles, Turner& Turner 80-26M* (TEX); 19mi [S] of Antiguo Morelos, Whiffin & Rodriguez 206 (TEX).-TAMAULIPAS:along Hwy 85, 111 km NW of Cd. Victoria, Croat 39215 (F, MO); 15 mi from Tampico on the Mante Hwy, Graham & Johnston 4089 (MICH, TEX); 3 mi E of Casas, Johnston 5776 (MEXU, MICH, TEX); El 1990 SIMSIA 57

Bordo, 19May 1966, Rzedowski s.n. (ENCB,WIS); N side of Tampico, Spooner2884* (MEXU, OS, P, RSA, WIS); along Tampico-Cd. Victoria Rd, 14.6 mi SE of jct of rd in town ofManuel to Cd. Mante, Spooner 2885* (OS, OSH, TENN, US); along Rt 180, 3.1 mi N of jct of rd toAldama, Spooner 2886* (CHAPA, F, MEXU, OS, WIS); 4 mi S of Los Lavaderos, Spooner 2887* (MEXU, OS); 8.1 mi NE of Manuel, Stuessy & Roberts 3635 (NY, OS, UC).-VERACRUZ: Jalapa-VeracruzRd, turnoff toLos Banios de Carrizal, Calzada 2052 (ENCB, F, LE, MEXU, NY), Spooner 2882* (MEXU, OS); 1 km N of Poza Rica, Hansen & Nee 7414 (MO); 10.8 mi SW of Panuco, Keil & Canne 9231* (OS); banks of Rio Plano, Rt 140, 39 km S of Jalapa, Long & Burch 3225 (F, NY); Idolos, Paray 2881 (ENCB); valley of the Rio Pinuco, 15mi SW of Tampico, Worth 23 (PH).

Simsia eurylepis is a distinctive species that has an overall resemblance to S. lagascaeformis. It differs from the latter by being self-compatible, in having anthers yellow throughout (without a purple distal band as in S. lagascaeformis), in preva lence of lobed leaves, and in having proximally winged petioles, sometimes with nodal discs. Blake (1913) distinguished this species from S. submollicoma largelyby the presence of winged petioles or discs, but these features vary within individual populations. Also variable ispeduncle length,with some inflorescenes tightly aggre gated, and others more open with peduncles to 10 cm long. Although this character is taxonomically useful in characterizing some species, as S. steyermarkii, it has no diagnostic value here. Simsia eurylepis is the only eradiate species of the genus, but it, like the other radiate species of Simsia is visited by diverse pollinators (bees, beetles, pers. obser.) and it may be partly outcrossing. The possible hybridization of S. eurylepiswith S. foetida to produce S. chaseae has been discussed earlier (seeArtifi cialHybridizations). Two anomalous specimens of possible relationship to S. eurylepis are notewor thy: Tamaulipas, vicinity of Tampico, Palmer 52 (BM), which has subequal phyllaries, rays, and black anthers; and Nuevo Leon, vicinity of Horsetail Falls, El Cercado, Duke 3992 (MO), which has harshly pubescent leaves, nearly subequal phyllaries, and brownish anthers. Neither population has been relocated in the field. More material is needed before proper affinities can be determined.

15. Simsia chaseae (Millspaugh) S. F. Blake, Proc. Amer. Acad. Arts 49: 385. 1913. Encelia chaseaeMillspaugh inMillspaugh & Chase, Publ. Field Columbian Mus., Bot. Ser. 3: 125. 1904.-TYPE: MEXICO. Yucatan: ruins of Kobah, 26 Nov 1865, Schott 911 (lectotype, designated by Blake, 1913: F-176020!; isolectotype: F!).

Taprooted annuals, but occasionally persisting and suffrutescent. Stems 0.3-3 m tall, 0.2-2.0 cm in diameter, erect, green to purple or yellow, glandular-puberulent and hispid. Leaves with petioles 1-7 cm long, 1-2 mm in diameter, without wings or discs; blades 3-18 cm long, 2-15 cm wide, broadly to narrowly ovate to deltate, herbaceous, unlobed to 3-lobed half way to the midrib, at base cordate to truncate or cuneate, apex acute to acuminate, glandular-puberulent, sometimes with thinly scat tered scabrous or hirsute vesture, margins crenate to dentate. Inflorescence corymbiform, loosely branched; peduncles 1-6.5 cm long, 1 mm in diameter, glandular-puberulent and hirsute. Heads radiate, 12-16 mm long, 10-15 mm in diameter. Involucre urceolate, 11-15 mm long, 7-12 mm in diameter; phyllaries 16 32, 3-seriate, unequal to almost subequal, appressed to slightly reflexed apically, green to yellow or brown, acute to acuminate at the tips, glandular-puberulent and 58 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 30 hispid, margins entire; outer phyllaries 5-9.5 mm long, 1.1-2 mm wide, ovate to lance-ovate; inner phyllaries 7.8-10.2 mm long, 1.2-2.2 mm wide, lanceolate. Pales 7.5-9.3 mm long, 1.8-3 mm wide, stramineous, sometimes flecked greenish black apically, acute or acuminate, scabrous apically. Ray florets 6-13; ligules 4.2-9 mm long, 1.5-3.2 mm wide, light lemon-yellow; tube 1-1.6 mm long, 0.3-0.5 mm in diameter; ovary 5-7.4 mm long, 0.4-0.5 mm wide. Disc florets 21-67; corolla 5.3 6.3 mm long, 0.8-1 mm in diameter; tube 0.8-1.6 mm long, 0.4-0.6 mm in diameter; lobes 0.4-0.8 mm long, 0.4-0.8 mm wide; anthers 2.2-3.2 mm long, yellow, rarely blackish distally, terminal appendages stramineous; achenes 4.2-6.1 mm long, 2.5 3.8 mm wide, laterally flattened, awns absent or 1.8-3.2 mm long, squamellae absent. Chromosome number: n = 17. Fig. 7. Phenology. Flowering from June throughNovember. Distribution (Fig. 19). Mexico, along the Gulf coastal lowlands in central and southern Veracruz, and in the northern part of the Yucatan Peninsula; roadsides, borders of agricultural fields, and other weedy areas; 0-600 m.

REPRESENTATIVE SPECIMENS. Mexico. CAMPECHE: near Balonchenticul, on Rt 261, 5.1 mi S of Campeche-Yucatan border, Spooner 2802* (MEXU, OS, OSH, TEX, UC, US).-QUINTANA Roo: Santa Rosa, jct of rd to Peto and rd to Tzucacab, Spooner 2811* (GH, MEXU, MO, OS, P, TEX, UC, US).-VERACRUZ: Cerro Gordo, Dorantes et al. 1743 (ENCB, MEXU, MO), Spooner 2880* (F, MEXU, OS, TENN, TEX, UC); Zacuapan, Purpus 7472 (UC), 11121* (F, NY, US), 13085 (MO), 13095 (F, K), 15002 (F); crossing of Jalapa-Huatusco Rd & Rio Pescado, Turner & Tapia 15522 (TEX); 13.4 mi W of Rt 140, near Puente Nacional, on rd to Huatusco, Spooner 2859* (OS); Jalapa-Veracruz Rd, at jct of rd to Carrizal, Spooner 2881* (MEXU, OS); 3.8 mi E of Tuzamapan, Zola 12 (TEX). YUCATAN:Uxmal, Degener 26757 (US), Gustafsson 1471 (US); Izamal, Gaumer 910 (CAS, F, G, GH, MO, NY, S); Chichankanab, Gaumer 2045* (B, BM, C, F, GH, MO, NY, S, US); Calotmul, Gaumer 2046 (CAS, LE, W); ruins of Kobah, Spooner 2803 (GH, LE, MEXU, OS); Rt 180, just NW of Santa Elena, Spooner 2804* (MEXU, OS); 2.7 mi SE of town square of Muna on rd to Ticul, Spooner 2805* (MEXU, OS); 4.9 mi S of town square of Uman on Rt 180, Spooner 2806 (OS); Progreso-Merida Rd, 4.4 mi S of rd to Yukalpeten, Spooner 2807a (OS); 6.1 mi NW of town square of Motul, on rd to Baca, Spooner 2807b (OS); 1.2 mi E of Temax, Spooner 2808 (OS); E of Buctzotz, on Motul-Tizimin Rd, 6.7 mi W of rd to Cenotillo, Spooner 2809* (OS); Chichen Itza, Steere 1026 (GH, LL, MICH, TEX), 1625 (MICH); Progreso, Steere 3027 (LL,MICH).

Simsia chaseae is remarkably similar to S. dombeyana, and it often is difficult to distinguish these species (see Numerical Taxonomy of S. chaseae and S. dom beyana). The former usually has outer phyllaries 3/4or less the length of the inner phyllaries, and the latter almost always has subequal phyllaries. In addition, S. chaseae tends to have smaller heads and fewer disc florets than S. dombeyana. The distribution (S. chaseae confined toMexico and S. dombeyana in South America) and phenology (S. chaseae flowering from June-November and S. dombeyana flow ering fromDecember-May) simplifiesmany identifications. Simsia chaseae is intermediate in many features between S. eurylepis and S. foetida, and possibly is of hybrid origin between them (see discussion under Artifi cial Hybridizations). Its overall resemblance is closer to S. foetida, with which it shares a yellowish color, highly glandular vesture, and spicy odor. It is best distin guished from this species by its sparingly glandular-pubescent leaves, which often appear almost glabrous to the naked eye, the usually shorter outer phyllaries lack ing a sinuate margin, the urceolate heads, and its self-compatibility. 1990 SIMSIA 59

16. Simsia dombeyana de Candolle, Prodr. 5: 578. 1836.-TYPE: .Lima, [1778-1788, fide Ruiz, 1940], Dombey 12 (holotype: P!, fragment: US!, photo: OS!; isotypes: B [fide Blake, 1917] destroyed, photos: CM! GH! TEX!; G-DC, microfiche: IDC. 800. 946. III. 8!, photos: GH! MO! NY! TEX!, tracing:GH!; MPU!). Encelia hirsutaKuntze, Revis. gen. pi. 3(2): 145. 1898. Simsia hirsuta (Kuntze) S. F. Blake, Proc. Amer. Acad. Arts 49: 389. 1913.-TYPE: ARGENTINA. Cordoba: Dique near Cordoba, Dec 1891, Kuntze s.n. (lectotype, here designated: US-701505!; isolectotypes: NY [2]!US!, fragment:GH!). Encelia hirsuta f. radiata Kuntze, Revis. gen. pl. 3(2): 145. 1898.-Type: PERU. Lima, [1778-1788, fide Ruiz, 1940], Dombey 12 (holotype B, [fideBlake, 1917] destroyed, photos: CM! GH! TEX!; isotypes: G-DC, microfiche: IDC. 800. 946. III. 8!, photos: GH! MO! NY! TEX!, tracing:GH!; MPU!; P!, fragment:US!, photo: OS!.

Taprooted annuals. Stems 0.4-3.5 m tall, 0.3-3.0 cm in diameter, erect, green to purple or yellow, glandular-puberulent and hispid. Leaves with petioles 1-8 cm long, 1-2 mm in diameter, without wings or discs; blades 3-15 cm long, 2-12 cm wide, ovate to deltate, herbaceous, usually unlobed but sometimes slightly lobed to lobed half way to the midrib, base cordate to truncate or cuneate, apex acute to acuminate, glandular-puberulent, with intermixed scabrous and hirsuite vesture, margins crenate to dentate. Inflorescence corymbiform, loosely branched; peduncles 2-15 cm long, 1-2 mm in diameter, glandular-puberulent and hirsute. Heads radiate, 12 18 mm long, 9-17 mm in diameter. Involucre urceolate, 10-16 mm long, 9-15 mm in diameter; phyllaries 16-32, 3-seriate, subequal, appressed to slightly reflexed at the tips, green to yellow or brown, lanceolate, acute or acuminate at the apex, glandular puberulent and hispid, margins entire; outer phyllaries 8-12 mm long, 1.6-3.1 mm wide; inner phyllaries 8-12.5 mm long, 1.2-2.9 mm wide. Pales 7.7-10.5 mm long, 1.8-2.8 mm wide, stramineous, sometimes purple or speckled black apically, acute to acuminate at the tips, scabrous apically. Ray florets 6-13; ligules 2.5-10 mm long, 1.3-2.9 mm wide, light lemon-yellow; tube 0.5-2.3 mm long, 0.2-0.5 mm in diame ter; ovary 3.7-8.2 mm long, 0.4-0.5 mm wide. Disc florets 23-82; corolla 4.2-8.4 mm long, 0.7-1.1 mm in diameter; tube 0.7-2.3 mm long, 0.4-0.6 mm in diameter; lobes 0.3-0.7 mm long, 0.4-0.6 mm wide; anthers 1.8-3.5 mm long, yellow to black, terminal appendages stramineous; achenes 5-7.5 mm long, 2.5-3.9 mm wide, later ally flattened, awns 1.2-4.6 mm long, squamellae rarely present. Chromosome num ber: n = 17. Figs. 6, 8, 11. Phenology. Flowering specimens can be found throughout the year, but most plants flower fromDecember throughMay. Distribution (Fig. 14). Scattered throughout the Andes from Ecuador and Peru to Bolivia, and the northern parts of Argentina, east to Entre Rios, Argentina, and disjunct in eastern in the states of Bahia and Minas Gerais; roadsides and disturbed ground; 40-2020 m.

REPRESENTATIVE SPECIMENS: Ecuador. GUAYAS: along rd to Daule, 17 km E of Palestina, King & Garvey 7008 (F, MO, NY, UC, US); 2 km N of Daule, Spooner 2907* (OS, TENN, UC) 3.5 km NE of Duran, Spooner 2908* (OS, TEX, US) .-LOJA: slopes of Cerro Villonaco, Harling & Andersson 13338* 60 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 30

(US). Peru. CAJAMARCA: 15 km E of Chilete on Cajamarca-Pacasmayo Rd, Dillon & Dillon 4618 (MO, OS); Km 124 of Pacasmayo-Cajamarca Rd, near Choropampa, Sdnchez 2368 (CHAPA, SI). LAMBAYEQUE:Pampa de Olmos, between Motupe & Piura, Ferreyra 12369 (MO).-LA LIBERTAD:Cerro Cabez6n, Sagdstegui & Cabanillas 8740 (F, SI).-LIMA: Km 56, Central Rd, Ferreyra 10482 (MO). PIURA:32 km S of Piura, Piura-Chiclayo Rd, Ferreyra 5902* (US); Hacienda Pabur, Piura-Huancabamba Rd, Ferreyra 6037* (F, NY, US); Cabo Blanco, Haught 249* (BM, GH, NY, US); between Piura & Nomala, Weberbauer 5934* (F, G, GH, US).-TUMBIs: Lechugal, Ferreyra 10662 (MO);Mancora Val ley, 50 mi N of Tumbds, Haught 155 (NY, US). Brazil. BAHIA:12 kmW of Irece, Coleman 505 (US); valley of Rio Formoso, 40 km SE of Correntina, Harley 21718 (K, US); 5 km N of Tobocas, Harley et al. 21987 (K,NY, US); 32 km S from rd between Itaberaba& Ibotirama, along rd toAndarai, King & Bishop 8704 (US).-MINAs GERAIS:2 km along rdN ofMato Verde towardsMonte Azul, King & Bishop 8586* (K, US). Argentina. CATAMARCA: Dept. Belen, 12.1 km W of Rt 40 on rd to Pozo de Piedra, E of La Toma, Spooner & Clausen 4604 (SI,WIS); between Alijilan & El Alzo, Crist6bal 328* (UC); Andalgana, Jorgensen 1286* (BAB, GH, MO, SI, UC), Turner 9232 (LL); Santa Maria, Morello 280* (LIL). C6RDOBA:near C6rdoba, Castellanos 94 (SI), Stuckert 6403 (SI); Pampas, Jan 1892, Kuntze s.n. (NY); Quebrada de Las Rosas, Lanfranchi 1314 (SI).-ENTRE Rios: Parana, Burkart 22337* (SI); Diamante, Burkart 22323 (SI).-Jujuy: Alto de la Vina, Cabrera & Kiesling 24908* (SI).-LA RIOJA:Dept. Chilecito, 12.2 km NW of Chilecito on rd to El Vallecito, Spooner & Clausen 4584 (SI,WIS).-SALTA: aeropuerto, Dept. Capital, Krapovickas & Schinini 35947 (OS); Prov. Cafayate, Rodriguez 1442* (BAB, SI).-SANTIAGODEL ESTERO: Dept. Guasayan, El Cevillar, Mar 1944, Pierotti s.n. (U).-TUCUMAN: Vipos, Dinelli 812 (BAB), Lillo 7951 (BAB), 25375 (GH, US), Venturi 4999 (US); San Pedro de Colalao, Venturi 6277* (US). Bolivia. COCHABAMBA:Mizque, Prov. Campero, Steinbach 711 (GH,MO, NY, UC, WIS).-TARIJA: Prov. Aviles, Chocloca, Beck & Sdenz 724 (LPB, SI, US).

Simsia dombeyana is distinguished by its subequal phyllaries, urceolate heads, unwinged petioles, and self-compatibility, all features possessed by some popula tions of S. chaseae. Simsia dombeyana lacks the spicy odors of S. chaseae.Although there is some overlap in relative inner vs. outer phyllary dimensions, most popula tions are readily distinguished (see Numerical Taxonomy). Simsia dombeyana showsmuch floralvariation. For example, Ferreyra 6037 has disc corollas 8.4 mm long with tubes 2.3 mm long, whereas in Burkart 22337 the measurements are 4.2 mm and 1.0 mm, respectively. These variants show no geo graphical correlation, however. One feature that does exhibit some geographic correlation is anther color, which is yellow in the northern part of the range and often black in the southern part (e.g., in the south Beck & Saenz 724, Cabrera & Kiesling 24908, Crist6bal 328, Dinelli 812, and Jorgensen 1286). One collection of uncertain relationship cited by Kuntze (1898) as a syntype of Encelia hirsuta f. radiata [Colombia. Bolivar: Cartagena, 1825-1826, Billberg s.n. (S, 4 specimens)] has sinuate phyllaries more typical of S. foetida but with an overall appearance of S. dombeyana. Burkart (1974) and Cabrera (1978) stated that many specimens of S. dom beyana are eradiate. Some specimens have very small rays that are not or only slightly exserted beyond the phyllaries, and the rays in this species, as in all others in the genus, are easily separated from the ovary; these features probably led Burkart and Cabrera to interpret the species as eradiate.

17. Simsia amplexicaulis (Cavanilles) Persoon, Syn. pl. 2: 478. 1807. Coreopsis amplexicaulis Cavanilles, Descr. pl. 226. 1802. Encelia amplexicaulis (Cavanilles) Hemsley, Biol. cent.-amer., Bot. 2: 183. 1881. Simsia am plexicaulis var. genuina, S. F. Blake, Contr. Gray Herb. 52: 46. 1917. TYPE: Cultivated in the Royal Botanic Garden of Madrid, from unvouch 1990 SIMSIA 61

ered achenes sent by the Sesse & Mocifio expedition from Mexico (holotype:MA!, photo: OS!). Ximenesia cordataHumboldt, Bonpland & Kunth, Nov. gen. sp. 4 [folio]: 178. 1818; 4 [quarto]: 228. 1820. Simsia cordata (Humboldt, Bonpland & Kunth) Cassini, Dict. sci. nat. 59: 137. 1829. Encelia cordata (Humboldt, Bonpland & Kunth) Hemsley, Biol. cent.-amer., Bot. 2: 183. 1881.-TYPE: MEXICO.Michoacain: "Valladolid de Mechoacan [Morelia, fide Sprague, 1924] et prope Capula Mexicanorum" [Aug-Sep 1803, fide Sprague, 1924], Humboldt & Bonpland 2350 (holotype: P-HBK, microfiche: IDC 6209. 109. I. 3!). Ximenesia heterophyllaHumboldt, Bonpland & Kunth, Nov. gen. sp. 4 [folio]: 178. t.380. 1818; 4 [quarto]: 227. 1820. Simsia kunthiana Cassini, Dict. sci. nat. 59: 137. 1829, nom. nov. Simsia heterophylla (Humboldt, Bonpland & Kunth) de Candolle, Prodr. 5: 577. 1836, non Simsia heterophylla (Cavanilles) Persoon, 1807. Encelia heterophylla (Humboldt, Bonpland & Kunth) Hemsley, Biol. cent.-amer., Bot. 2: 184. 1881.-TYPE: MEXICO. Michoacain: near "Zinapequaro" [Zinapecuaro], "alt 960 hex" [5808 ft, Sep-Oct 1803, fide Sprague, 1924], Bonpland 4360 (holotype: P-HBK, microfiche: IDC 6209. 109. I. 2!; isotype: P!). Helianthus trilobatusLink, Enum. hort. berol. alt. 2: 352. 1822.-TYPE: Culti vated in the Botanic Garden at Berlin, from achenes sent fromMexico. (holotype: B, destroyed). Simsia auriculata de Candolle, Prodr. 5: 577. 1836.-TYPE: MEXICO,near Mex ico City, Sep 1827, Berlandier 927 (lectotype, here designated: G-DC, microfiche: IDC 800. 946. III. 4!; isolectotypes: BM! F! Fl! G[3]! GH! LE! MO! OXF! P!W[3]!). Encelia mexicana Martius ex de Candolle, Prodr. 5: 578. 1836, pro syn. Ximenesia hirtaMartius ex de Candolle, Prodr. 5: 578. 1836, pro syn. Helianthus amplexicaulis de Candolle, Prodr. 5: 589. 1836.-TYPE: MEXICO. Guanajuato: Province of Leon, W of the town of "Guanaxuato" [Guana juato], 1829, Mendez s.n. (holotype: G-DC, microfiche: IDC 800. 951. I. 6!, fragment:GH!). Simsia schaffneri Schultz-Bipontinus ex A. Gray, Proc. Amer. Acad. Arts 19: 8. 1883, pro syn. Helianthus sericeus Sesse & Mocifio, P1. nov. Hisp. 147. 1890.-TYPE: MEXICO. [Morelos]: "Ayacapixtla" [Yecapixtla], "Sep" [probably Nov 1788, fide McVaugh, 1977], Sesse & Mocifio s.n. (holotype: MA; probable isotype: OXF!, photo: OS!). Simsia foetida var. decipiens S. F. Blake, Proc. Amer. Acad. Arts 49: 387. 1913. Simsia amplexicaulis var. decipiens (S. F. Blake) S. F. Blake, Contr. Gray Herb 52: 46. 1917.-TYPE: MEXICO.Chihuahua: "SW Chihuahua, Aug-Nov 1885" [Norogachic, 13-25 Nov, fideMcVaugh, 1956], Palmer 440 (holotype: GH!; isotypes: K! LE! LL!MEXU! MO! NY[2]! UC!).

Taprooted annuals. Stems 0.1-3 m tall, 0.1-1.0 cm in diameter, ascending or erect, sometimes decumbent at the base and rooting at the nodes, green to yellow, brown, or purple, glandular-puberulent and scabrous to hispid with intermixed hirsute vesture. Leaves with petioles 1-11 cm long, 1-2 mm in diameter, often 62 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 30 winged throughout and clasping at the dilated base, or only partially winged, or entirely unwinged; blades 2-18 cm long, 1-14 cm wide, ovate to deltate, herba ceous, narrowly to deeply 3-lobed or unlobed, base cordate to truncate or cuneate, apex acute to acuminate, scabrous and strigose, sometimes hirsute, usually egland ular, margins serrulate to dentate. Inflorescence corymbiform to paniculate, loosely branched; peduncles 1-12 cm long, 1-1.5 mm in diameter, glandular puberulent, scabrous and hirsute. Heads radiate, 12-14 mm long, 22-35 mm in diameter. Involucre ovoid-campanulate, 10-12 mm long, 7-13 mm in diameter; phyllaries 15-22, 2-3-seriate, usually nearly subequal, with the outer phyllaries usually slightly shorter than the inner, moderately to greatly reflexed, acute to rarely caudate at the tips, dark green to black, rarely purple, scabrous and hirsute, sometimes glandular, margins entire; outer phyllaries 5-14 mm long, 1-2.5 mm wide, lanceolate; inner phyllaries 6.1-14 mm long, 1.3-2.2 mm wide, lanceolate to linear. Pales 6.7-10.2 mm long, 1.2-3.2 mm wide, stramineous, usually flecked with black, especially above, rarely purple, acute to acuminate, antrorsely pubes cent apically. Ray florets 8-14; ligules 8-16 mm long, 3-7.5 mm wide, orange yellow; tube 0.7-1.7 mm long, 0.3-0.5 mm in diameter; ovary 4.6-6.5 mm long, 0.5-0.9 mm wide. Disc florets 23-55; corolla 5.2-7 mm long, 1-1.5 mm in diame ter; tube 0.8-1.3 mm long, 0.5-0.7 mm in diameter; lobes 0.8-1.3 mm long, 0.6 0.8 mm wide; anthers 3.2-4.2 mm long, yellow proximally, usually purple (often drying bronze) distally, with terminal appendages stramineous; achenes 3.5-5.5 mm long, 1.8-3 mm wide, laterally flattened, awns 2.2-3.8 mm long or more rarely absent, squamellae absent. Chromosome number: n = 17. Phenology. Flowering throughout the year, butmostly fromSeptember through November. Distribution (Fig. 20). Mexico, northern Chihuahua to Nuevo Leon, south to Honduras; mostly in openings in regions of oak-pine forests, along roadsides, irriga tion ditches, agricultural fields, wet upland meadows; 1300-2990 m.

REPRESENTATIVE SPECIMENS. Mexico. : Hwy 70, 3 mi E of La Sauz, Sundberg & Lavin 2922 (OS, TEX).-CHIAPAS: N edge of San Crist6bal de las Casas, Breedlove & Raven 12434 (CAS, DUKE, MEXU, MICH), 13283 (DS, ENCB, F, LL, MICH, NY, US); Tuxtla Gutierrez, Spooner 2780 (MEXU, OS); Rt. 195, 0.6 mi E of footpath to Pinabeto, Spooner 2794 (COL,MEXU, OS, UC, US); rd to Zinacantan, 2 mi N of San Crist6bal de las Casas, Stuessy & Gardner 4304 (ASU, CHAPA, ENCB, MO, OS, WIS).-CHIHUAHUA: ?2mi W of Cuauhtemoc, Powell & Edmondson 998 (TEX); 15mi SW of Buenaventura, Spooner & Schilling 2453 (BM, COL, G, GH, LL, MEXU, MICH, NY, OS, OSH, P, TEX, UC, US); E of Cuauhtemoc, Spooner & Schilling 2455 (LE,MEXU, MICH, NY, OS, P, UC, US).-COAHUILA: Saltillo, Palmer 422 (BM, C, F, G, GH, MSC, MO, NY, S, US); Parras, Palmer 427 (BM, C, F, G, GH, MEXU, MO, MSC, NY, UC, US); Hwy 57, 6 mi SE of turnoff toArteaga, Spooner & Schilling 2463 (BM, COL, G, GH, LE, MEXU, MICH, NY, OS, TEX, UC). DISTRITOFEDERAL: near Mexico City, Berlandier 850 (BM, G, LE, OXF, W); Xochimilco, Rzedowski 1062 (CHAPA, ENCB, IJ, MEXU, OS).-DURANGO: Durango, Palmer 657 (C, CAS, F, G, GH, MEXU, MICH, MO, NY, UC, US); Rt 45, 19.7 mi N of traffic circle on N side of Durango, Spooner & Brandenburg 2326 (CHAPA, OS).-GUANAJUATO:"Guanaxuato," Hartweg 145 (BM, CGE, G, OXF, P); 1 km W of San Luis de La Paz, Garcia 64 (CHAPA, MEXU).-HIDALGO: Rt 105, 10 mi SE of Pachuca, Spooner & Brandenburg 2229 (CHAPA, OS); Mpio. de Tepeapulco, Rancheria Los Cides, Ventura 1723 (ARIZ, CAS, CHAPA, ENCB, MEXU, OS).-JALISCO: fields near Guadalajara, Pringle 4266 (BM, BR, F, G, GH, GOET, JE, LE, M, MASS, MEXU, MO, MSC, NY, S, TEX, UC, US, W); Rt 80, bypass around Tepatitlan, Spooner & Brandenburg 2302 (CHAPA, OS); Rt 80, W limits of Jalostotitlan, Stuessy & Gardner 4095 (ASU, MO, OS, WIS); Rt 80, between Lagos de Moreno & Ojuelos, Stuessy & Gardner 4083 (ASU, MO, OS).-MExIco: Rt 85, 20 mi N of rd to pyramids (Rt 30), 1990 SIMSIA 63

I ______r______960 goo

TroplicoJCancer

2O~~~~~~~~~~~~~~~~~~~~~2

4~~~~~~~~~~~~~~~~~~~~~~

0S. amplexicaulis

960 goo

FIG. 20. Distribution of Simsia amplexicaulis.

Powell & Edmondson 599 (MICH, TEX), 603 (MICH, TEX); 1mi W of Tepetlaoxtoc, on rd to Jalopan, Spooner & Brandenburg 2232 (CHAPA, OS); Rt 15, 7 mi W of , Stuessy & Gardner 4192 (OS); Rt 55, 0.5 mi S of Tenancingo, Stuessy & Gardner 4196 (ASU, ENCB, MO, OS, WIS).-MICHOACAN: Rt 15, 14 mi W of Zamora, Powell & Edmondson 840 (MICH, TEX); Rt 15, near intersection of rd to Zinapecuaro, Spooner & Villasefior 2557 (MEXU, OS, OSH, US).-MORELOS: Coajomulco, Diaz 153 (DS, ENCB, MICH, NY). -NUEvo LE6N: 8.7 mi NW of San Rafael, Sundberg et al. 1744 (OS, TEX). OAXACA: Rt 190, 1 mi SE of Asunci6n Nochixtlan, Spooner & Dorado 2654 (B, G, LE, MEXU, MO, OS, UC, US); Rt 190, 10 mi SE of Nochixtlan, Stuessy & Gardner 4259 (OS).-PUEBLA: Rt 190, just E of San Martin Texmelucan, King 2239 (TEX, US); Rt 140, 22 mi NE of Tepeaca, Powell & Edmondson 640 (F, MEX, MICH); 3 mi SE of Cd. Serdan, Spooner & Triplehorn 2048 (CHAPA, OS, UC, US, WIS).-QUERETARO: Cadereyta, Kelly 672 (UC).-SAN Luis POTOSI: 2 mi S of Venado, Spooner & Brandenburg 2195 (OS).-TLAXCALA: San Juan Quetzalcuapan, Vibrans 410 (ARIZ, MEXU, OS, U).-VERACRUZ: Guadalupe Victoria, Turner 15197 (TEX).-: 8 mi E of Zacatecas on rd to Aguascalientes, Garcia 805 (CAS, CHAPA, TEX); Rt 85, 10 mi SE of Arriaga, Spooner & Villasenior 2488 (OS). Guatemala. BAJA VERAPAZ; Purulha, Spooner & Dorado 2751 (MEXU, OS, TENN, UC).-CHIMALTENANGO: Aldea El Parcelamiento, near Chocoyos, Spooner & Dorado 2684 (MEXU, OS, UC).-EL QUICHE: Utatlan Ruins, Molina & Molina 25095 (C, F, NY, US). -HUEHUETENANGO: Rt 9N, 3 mi S of Huehuetenango, King 3415 (DS, MICH, NY, TEX, UC).-JALAPA: between Jalapa & Montania Miramundo, Steyermark 32894 (F, GH).-QUEZALTENANGO: vicinity of Zunil, Standley 83164 64 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 30

(F).-SACATEPtQUEZ:Volcan de Agua near SantaMaria de Jesus, Spooner & Dorado 2687 (F,MEXU, MO, OS).-SAN MARCOS:near Castalia, Spooner & Dorado 2673 (MEXU, OS).-SOLOLA: near Nahuala, Williams et al. 41437 (F, NY).-TOTONICAPAN:jct of Rt CA-1 & rd to San Francisco el Alto, Stuessy & Gardner 4335 (ENCB, OS). Honduras. MORAZAN:Cerro de Uyuca, Sauer 1582 (UC,WIS).

Simsia amplexicaulis is distinguished by its annual habit, unwinged petioles, nodes lacking nodal discs, leaves with no or few glands, subequal phyllaries, and anthers that are yellow proximally and usually purple distally. It sometimes is difficult to distinguish herbarium specimens of Simsia am plexicaulis from S. foetida var. foetida. Both have subequal to nearly subequal phyllaries, often with caudate tips. Simsia amplexicaulis generally has green, non glandular leaves and dark green to blackish phyllaries; S. foetida var. foetida gener ally has yellowish, densely stipitate-glandular leaves and yellowish phyllaries. These differences are very pronounced in the field, where the glandular, spicy smelling and viscid leaves of S. foetida are easily distinguished from those of S. amplexicaulis. Additionally, these species grow in very different habitats. Although there is some overlap in their absolute elevational ranges, S. foetida grows in the hot lowlands and S. amplexicaulis grows in the cool uplands; they rarely co-occur. Simsia amplexicaulis is an extremely variable species. Some plants may be profusely branched and attain a height of 3 meters, whereas others within the same population may be stunted, with unbranched individuals flowering at 0.1 meter tall. Leaf shape is especially variable, with petiolar wings absent or present, and the lamina lobed or entire. The species is one of the dominant weeds in upland areas in Mexico (Rzedowski 1983) and is frequently found along the borders of agricultural fields, along irrigation ditches, about human dwellings, and even growing from cracks in asphalt along roads. In less disturbed settings, it is a component of pine-oak forests. The type specimen for Simsia amplexicaulis was grown from seed from the Sesse and Mocinlo expedition, probably sent by Vincente Cervantes (Cavanilles 1791; McVaugh 1977). No specific locality is provided by Cavanilles, and this expedition covered much of central, western, and southern Mexico (McVaugh 1977). A mixed sheet bearing four fragmentary specimens, labelled Coreopsis am plexicaulis and C. foetida in Cavanilles's handwriting (comparison from Burdet, 1973) represents a possible isotype. The holotype of S. amplexicaulis var. decipiens has the calvous achenes that were used to distinguish this variety, but the isotype atUS bears awned achenes, an intrapopulational variation pattern common within many populations of various taxa of Simsia.

18. Simsia foetida (Cavanilles) S. F. Blake, Proc. Amer. Acad. Arts 49: 385. 1913. Coreopsis foetida Cavanilles, Icon. 1: 55. t. 77. 1791. Simsia ficifolia Persoon, 1807, nom. superfl. Ximenesia foetida (Cavanilles) Sprengel, Syst. veg. 3: 606. 1826. Encelia foetida (Cavanilles) Hemsley, Biol. cent. amer., Bot. 2: 183. 1881.-TYPE: Cultivated in the Royal Botanic Garden ofMadrid, from unvouchered achenes sent by the Sesse & Mocifio expedi tion (holotype: MA!, photo: OS!; isotype:MA!).

Taprooted annuals, occasionally persisting and woody at base. Stems 0.5-4 m tall, 2-20 mm in diameter, erect, yellow-green to green or purple, usually copiously 1990 SIMSIA 65

glandular-puberulent, setose or pilose, sometimes hispid. Leaves with petioles 1 10.5 cm long, 0.5-1.5 mm in diameter, unwinged or winged, sometimes auriculate clasping at base with stipules of opposite leaves absent or present, generally un fused, very rarely fused with those of opposite leaves to form nodal discs; blades 3 18 cm long, 2-16 cm wide, ovate to deltate, herbaceous, unlobed, or narrowly to more rarely deeply 3-lobed, base cordate to truncate or cuneate, apex acute to acuminate, hirsute to sericeous, intermixed glandular-puberulentwith stipitate or subsessile glands, margins crenate-dentate. Inflorescence of solitary heads or of loosely and sparingly branched peduncles arising frommany nodes; peduncles 1-14 cm long, 1-2 mm in diameter, densely glandular-puberulent and hirsute. Heads radiate, 10-20 mm long, 20-45 mm in diameter. Involucre campanulate to ovoid campanulate, 8-17 mm long, 8-22 mm in diameter; phyllaries 11-66, 2-4-seriate, subequal, reflexed throughout to appressed proximally and reflexed only at the very tips, yellow-green to light-green, linear to linear-ovate, usually sinuate, caudate to rarely acute at the tips, hirsute or velutinous and densely glandular puberulent, with a single sinuate constriction near the apex; outer phyllaries 6-20 mm long, 1.2-3.5 mm wide; inner phyllaries 7.1-20 mm long, 0.5-3.2 mm wide. Pales 6.5-14 mm long, 1.3-3.8 mm wide, stramineous, acute to attenuate, scabrous apically. Ray florets 7-45; ligules 3.8-12 mm long, 1.5-7 mm wide, light lemon yellow to orange-yellow; tube 1.1-2.1 mm long, 0.3-0.9 mm in diameter; ovary 2.3-9.5 mm long, 0.7-1 mm wide. Disc florets 20-172; corolla 5.2-8.7 mm long, 1 1.5 mm in diameter; tube 0.6-2.5 mm long, 0.5-0.8 mm in diameter; lobes 0.5-1.5 mm long, 0.7-1 mm wide; anthers 2.7-4.5 mm long, yellow or rarely black, termi nal appendages stramineous; achenes 3.1-7.5 mm long, 1.7-4.1 mm wide, laterally flattened, awns 1.7-7.4 mm long, squamellae rarely present. Chromosome num ber: n = 17 (unknown for var. jamaicensis). Simsia foetida is easily distinguished by its densely glandular vesture and subequal phyllaries, often with caudate tips and a single sinuate constriction near the apex. It is separated from the very similar S. chaseae and S. dombeyana by its campanulate to ovoid heads (urceolate in S. chaseae and S. dombeyana). It is easy to confuse some herbarium specimens of S. foetida with S. amplexicaulis, although in living condition the highly glandular, spicy-smelling leaves of S. foetida easily distinguish it from S. amplexicaulis, which has no or few leaf glands and no spicy odor. Simsia foetida has yellow anthers; those of S. amplexicaulis are yellow proxi mally and have a purple (drying bronze) band distally. Simsia foetida is a highly variable species here interpreted to comprise five varieties distributed in Mexico from Jalisco to Queretaro, then south to Panama, with a disjunct variety in Jamaica and disjunct populations in Bolivia and Venezu ela. These varieties differ mainly by overlapping quantitative features, but previous treatments have ignored this overlap and have described these differences in a qualitative fashion or in a more discrete quantitative way. Some intergradation of these varieties sometimes makes identifications difficult. For a fuller discussion of these problems, see the Numerical Taxonomy section and the discussions below.

18a. Simsia foetida var. foetida. Encelia adenophora Greenman, Proc. Amer. Acad. Arts 39: 109. 1903. Simsia adenophora (Greenman) S. F. Blake, Proc. Amer. Acad. Arts 49: 388. 1913.-TYPE: MEXICO. Jalisco: Tequila, fields and copses, Sep-Oct 1893, 66 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 30

Pringle 4602 (lectotype, designated by Blake, 1913: GH!; isolectotypes: BM! BR! F! G[21! GOET! JE! K! LE! M! MEXU[2]! MSC! NY! S! UC! US[2]!). Simsia guatemalensis H. Robinson & R. D. Brettell, Phytologia 24: 372. 1972.-TYPE: GUATEMALA.Guatemala: Amatitlan, 1200 m, Oct 1904, Tuerckheim 7801 (holotype: US!, photo: OS!).

Stems 1-4 m tall. Leaves with petioles 1-10.5 cm long, unwinged or narrowly winged, sometimes auriculate-clasping, at base stipules 0-2 cm long, sometimes united with those of opposite leaves to form nodal discs; blades 3-11.5 cm long, 2 11 cm wide, densely glandular-puberulent with stipitate glands, setose, sometimes hispid. Peduncles 1-9.5 cm long. Heads 11-16 mm long, 20-30 mm in diameter. Involucre 8-17 mm long, 8-13 mm in diameter; phyllaries 13-26, 2-3-seriate, hirsute and glandular-puberulent; outer phyllaries 6-14 mm long, 1.2-2.8 mm wide; inner phyllaries 7-14.5 mm long, 1.2-3.2 mm wide. Pales 7-12 mm long, 1.5-3.8 mm wide. Ray florets 7-14; ligules light lemon-yellow, 5.5-11 mm long, 2.8-4.6 mm wide; ovary 3.4-9.5 mm long. Disc florets 20-76; corolla 5.2-8.2 mm long, tube 0.6-1.8 mm long; achenes 3.5-6.6 mm long, 1.8-4 mm in diameter, awns 1.7-5.6 mm long. Chromosome number: n = 17. Fig. 9. Phenology. Flowering from July throughMarch. Distribution (Fig. 21). Southern Nayarit east to Queretaro, southeast to El Salvador; on open grassy fields, dry weedy thickets, streamsides, or along road sides; in areas of tropical deciduous forests; 300-2150 m, but rare above 1700 m.

REPRESENTATIVESPECIMENS. Mexico. CHIAPAS:Mpio. of Tenejapa, paraje of SibanilHa', near school house of Pokolum, Breedlove 9437 (DS, ENCB, LL, MICH, NY); atMirador of El Sumidero, Breedlove 41483 (DS, MO, TEX), Spooner2787* (MEXU, OS); Rt 190, 0.3 mi S of San Gregorio, Spooner2768* (MEXU, OS, P, TEX, UC, US); rd into Parque Natural Ejidal Lagos de Col6n, 3 mi E of Rt 190, just N of San Gregorio, Spooner2769* (BM, COL, LE, MEXU, MICH, OS, OSH, TENN, UC, US, WIS); Hwy 190, 4.6 mi S of rd into La Trinitaria, Spooner 2770 (MEXU, OS); pull-off area for Mirador El Roblar on rd toEl Sumidero fromTuxtla Gutierrez, Spooner2786* (MEXU, OS); N of Arriaga toTuxtla Gutierrez, 1.9 mi S of jct to Santa Elena, Spooner 2792* (F, GH, MEXU, MICH, NY, TEX); Mpio of Tenejapa, paraje of Mahbenchauk, barrio of Tih Ha', Ton 2033 (DS, ENCB, F, MICH, MSC, NY, US). GUANAJUATO:23 mi N of Celaya on rd toQueretaro, Crutchfield& Johnston 5951A (TEX).-GUERRERO: Coyuca, Hinton 5538* (BM, G, K, US); Cafi6n de Zopilote, 4 km N of Milpillas, Rzedowski 23516* (CHAPA, ENCB, MEXU, OS); Rt 95 toll rdon N side of Iguala, Spooner2846* (MEXU, OS).-JALISCO: Rt 80 to Guadalajara, 0.5 mi E of jct of rd to Acatlan, Spooner & Brandenburg2292 (MEXU, OS, US); Rt 80, 37mi SW of Guadalajara, Spooner & Brandenburg2300 (MEXU, OS, UC); S limitsof Atenquique on short rt toGuadalajara, Stuessy & Gardner3088 (F,MEXU, MSC, NY, OS, UC); 26mi NW of Tequila on Rt 15, Stuessy & Gardner 4111* (ASU, ENCB, MO, OS); 2 mi N of Guadalajara on rd to Saltillo, Villarreal 1264* (MICH).-MtxIco: Rincon del Carmen, Hinton 1741* (BM, G, GH, K, MEXU, NY, US); Volcan,Hinton2216* (F,GH, K, LL, MO, NY, PH); Acatitlan, Hinton8582* (K,LL, GH, K);6km N of Huehuetoca, Rzedowski 29500* (ARIZ, ASU, CAS, COL, ENCB, US); Rt 130, 4 mi SW of Tejupilco de Hidalgo, Spooner & Brandenburg 2264 (MEXU, OS); NW of , on Valle de Bravo-Tingambato Rd, 1mi NE of rd to Zacazonapan, Spooner & Villasenior 2569 (F, MEXU, MICH, OS, TENN), Villasefior & Spooner 772 (MEXU, OS).-MICHOACAN: near Barranca Hondo, 47 km S of Uruapan, Cronquist & Fay 10816* (CAS, DUKE, ENCB, F, MEXU, MICH, MSC, NY, RSA, TEX, US); Zitacuaro-Los Guajes, Hinton 13289* (ARIZ, ENCB, GH, K, LL, MICH, TEX, US).-MORELOS: Xochitepec, Lyonet 845* (MEXU, LL, US); limestone hills near Yautepec, Pringle 9898* (CM, F, GH, MO, NY, US).-NAYARIT: Ahuacatlan, Gentry et al. 19532* (DES, LL, US).-OAXACA: Monte Alban ruins,Smith 236 (MO, NY, TEX, US), Solano 384 (TEX), Spooner2822* (MEXU, OS, TEX); Rt 195, 61 mi SE of Oaxaca, Spooner & Dorado 2664* (COL, MEXU, NY, OS, P, US); Rt 195, 3 mi SW of San Jose 1990 SIMSIA 67

960 0oo

*. .* ;. . . 204

Smsiafoetid * var. foetida A var. megacephala -t * var. grandifl!ora * var.panamensis ,f *i: var. Jamaicensis ;Cw

96? 9Q0

FIG. 21. Distribution of the five varieties of Simsia foetida. See Fig. 14 for the distribution of of S. foetida var. panamensis inVenezuela and Bolivia.

Viejo, Spooner & Dorado 2666* (MEXU, OS, UC); 1mi N of Jayacatlan, Spooner2829* (MEXU, OS); 1 mi N of Monte Alban Park on rd to Cd. Oaxaca, Stuessy & Gardner 4261 (OS, WIS).-PUEBLA: Rt 190, 3 mi N of Tehuitzingo, Olsen & Lane 389 (LL), Spooner & Dorado 2637* (MEXU, OS, TEX). QUERETARO: Alamos, Arguielles 271* (ENCB, MEXU, NY). Guatemala. BAJA VERAPAZ: 4 mi E of town square of Salama, Spooner & Dorado 2747 (F, MEXU, MICH, OS, UC); near town square of San Miguel Chicaj, Spooner & Dorado 2749* (MEXU, OS, UC, US).-CHIQUIMULA: Rt 20, lower and middle slopes of Cerro Sill6n, 5 mi N of Guatemala-Honduras border, Stuessy & Gardner 4377 (OS). EL PROGRESO: Guatemala-El Progreso Rd (Rt 4), near border with Guatemala state, 8.4 mi NE of railway station at Agua Caliente, Spooner & Dorado 2691 (GH, LE, MEXU, OS, TENN, WIS); Guatemala City-Puerto Barrios Rd, 0.9 mi NE of intersection to Sanarate, Spooner & Linder 2758 (MEXU, OS).-EscUINTLA: near El Llano, Harmon 4668 (ENCB).-GUATEMALA: near Amatitlan, Holway 624* (GH), Standley 61469* (F); along Guatemala City-Puerto Barrios Rd, by bridge over river, 5.4 mi SW of Agua Caliente, Spooner & Dorado 2688* (F, GH, MEXU, OS, OSH, TENN, UC, US); 1.4 mi SW of S entrance to Amatitlan, Spooner & Dorado 2693* (BM, COL, MEXU, MICH, NY, OS, OSH, RSA, UC, US, WIS).-HUEHUETENANGO: mountains near El Reposo, 8 km from Mexican bor der, Williams et al. 41109 (MICH, F), 41366 (MICH, F).-JALAPA: vicinity of Jalapa, Standley 46514 (F). -SANTA ROSA: Cerro Redondo, Heyde & Lux 6160 (BR, F, G, GH, MO, NY, US). El Salvador. CHALATENANGO: along Tejutla Creek, hwy to La Palma, Molina & Montalvo 21575* (F, NY, US). Putative hybrids between S. foetida var. foetida and S. amplexicaulis: Mexico. HIDALGO: Mpio of Huehuetoca, W side of Cerro Sincoque, Rzedowski 34445 (CAS, CHAPA, ENCB, MEXU, NY, OS); just E of Mexico-Hidalgo state line on toll Rd 57 to Queretaro, Stuessy & Gardner 4578 (ASU, ENCB, MO, OS, WIS).-JALISCO: 48 mi S of Guadalajara, Cronquist 10290 (ENCB, MEXU, MICH, NY); Rt 15, 11 mi W of Jalisco-Michoacan state line, Lane & Olsen 2144 (TEX); 15 km SSE of Acatlan de Juarez, McVaugh & Koelz 341 (ENCB, MICH); Km 18-9 of rd to Sahuayo-Tizapan, Panero & Schilling 459 (OS, TENN); 2 mi W of S. L. Soyotlan, Spooner & Villasefior 2535 (BM, MEXU, MICH, OS, TENN, TEX, UC).-MICHOACAN: km 543, 8 mi NW of Sahuayo, McVaugh 18185 (DUKE, ENCB, MICH, US); rd to Jiquilpan de Juarez, 8 mi SE of Tizapan El Alto, Spooner & Villasenior 2538 (MEXU, OS). Putative hybrids between S. foetida var. foetida and S. lagascaeformis: Mexico. OAXACA: along rd from Huajuapan de Le6n to Mariscala de Juarez, via Amatitlan, 26.2 mi from the intersection of rd to San Agustin Atenango, Spooner & Dorado 2644 (BM, COL, F, LE, MEXU, MICH, NY, OS, OSH, 68 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 30

RSA, TENN, TEX, UC, US, WIS), same but 29.4 mi S, Spooner 2645 (OS).-PUEBLA:Km 170, 6.4 mi NE of center of Tehuitzingo on rd to Izucar, Spooner & Dorado 2635 (BM,MEXU, OS, TENN, TEX, UC, US); 12 mi SE of Tehuitzingo on rd to Oaxaca, Spooner & Dorado 2639 (MEXU, MICH, OS, TEX, UC).

All populations of S. foetida var. foetida from Nayarit to Oaxaca (northern populations) have petioles without stipules (except one collection fromQueretaro, Arguelles 271). The majority of populations from Chiapas, Mexico, to Guatemala (southern populations) have stipules and nodal discs, but there is considerable intrapopulational variation for this feature, with some nodes or entire individuals within populations lacking the structures. The results of the numerical analyses (see Numerical Taxonomy) suggest recognition of the northern and the southern popula tions as the same taxon. Nash (1976) considered S. foetida var. foetida (as S. foetida) and S. foetida var. grandiflora (as S. grandiflora) both to occur inGuatemala, but distinguished these taxa on the basis of presence/absence of stipules and nodal discs as well as size of the heads. My studies indicate that stipulemorphology is not a reliable discrimina tor between these taxa. In central and eastern Guatemala, some populations of these two taxa intergrade and are difficult to identify. Apparent hybrid swarms between S. foetida var. foetida and S. lagascaeformis are found with the co-occurring parents and are listed above. These putative hy brids are quite variable, exhibiting various combinations of parental features. Some have unequal phyllaries tinged with purple, as in S. lagascaeformis; all have the glandular leaves and lemon-yellow ligules, as in S. foetida var. foetida. Simsia foetida var. foetida apparently also hybridizes with S. amplexicaulis. Many of these hybrids occur about Lake Chapala in Jalisco and adjacent Michoacan andwere noted byMcVaugh (1984). Other apparent hybrids between S. foetida var. foetida and S. amplexicaulis occur in Hidalgo. These hybrids, like those with S. lagascaeformis, have densely glandular leaves like S. foetida var. foetida, yet possess heads like S. amplexicaulis with darker phyllaries and orange-yellow ligules. The Lake Chapala populations are all found between 1480-1650 m elevation, which is transitionalbetween the elevational ranges of the parents. The Hidalgo populations occur at higher elevations, from 2250-2500 m. Blake (1913) confused this taxon with S. amplexicaulis, which he cited under the name of S. foetida, and included S. foetida var. foetida as circumscribed here in S. adenophora. Blake (1917) later reported having seen an "authentic specimen" of Coreopsis foetida at G-DC and corrected his error.

18b. Simsia foetida var. jamaicensis (S. F. Blake) D. M. Spooner, comb. et stat. nov. Simsia jamaicensis S. F. Blake, Proc. Amer. Acad. Arts 49: 388. 1913.-TYPE: JAMAICA.Vicinity of Kingston, 500 ft, 29 Jan-4 Feb 1900, Clute 2 (holotype: GH!; isotypes: F! K! MO! NY! US[21!, fragment:G!).

Stems 1-3 m tall. Leaves with petioles 1-5.5 cm long, unwinged, blades 3-17 cm long, 2-16 cm wide, densely glandular-puberulent with subsessile glands, setose, sometimes hispid. Peduncles 1-6 cm long. Heads 10-18 mm long, 20-30 mm in diameter. Involucre 8-17 mm long, 8-12 mm in diameter; phyllaries 11-18, 2-3-seriate, hirsute and glandular-puberulent; outer phyllaries 6.5-11.4 mm long, 1990 SIMSIA 69

1.7-2.5 mm wide; inner phyllaries 8-10.8 mm long, 1.6-2.2 mm wide. Pales 8.2 10.6 mm long, 2-2.6 mm wide. Ray florets 7-10; ligules 3.8-6.8 mm long, 1.5-3.8 mm wide, light lemon-yellow; ovary 6.5-8.7 mm long. Disc florets 25-54, corolla 6.3-7 mm long, tube 1.7-1.9 mm long; achenes 5.8-7.5 mm long, 2.5-3.5 mm wide, awns 3.3-5.4 mm long. Chromosome number unknown. Phenology. Flowering fromNovember throughFebruary. Distribution (Fig. 21). Known only from within a 20 km radius of Kingston harbor, Jamaica; open dry limestone slopes; 30-200 m.

ADDITIONAL SPECIMENS EXAMINED. Jamaica. MIDDLESEX: St. Catherine Parish: upper slopes of Port Henderson Hill, Proctor 32765* (ENCB, IJ, LL, MASS, NY, RSA); Rodney's Lookout near Port Henderson, Wunderlin 5081* (NY, F).- SURREY:Kingston Parish: base of Long Mountain behind Rennock Lodge, Proctor 24496* (GH, IJ, LL, MICH, NY, US). St. Andrew Parish: valley of Hope River, University Rd toHope Rd, Adams 8766* (BM, DUKE, M); Kings House Grounds, Harris 6953 (NY, F), 6989* (B, BM, F, K); Hope Grounds, Harris 8228* (BM, F, NY); Long Mountain Road, Harris 10001* (BM, C, F, G, NY, US); 1mi SSE of Papine, Proctor 21945* (IJ, LL). St. Thomas Parish: Four-mile wood near milepost 13, Proctor 38472* (IJ); on Hwy A-4, 2 mi W of Albion, Pruski 1269* (IJ, K, NY, OS, US).

Simsia foetida var. jamaicensis appears to combine some features of var. foetida and var. grandiflora. It has the smaller heads and lemon-yellow ligules of var. foetida, and the darker green-yellow leaves with subsessile glands of var. grandi flora. It has disc floretswith consistently larger tubes than other varieties. The occurrence of all sites of S. foetida var. jamaicensiswithin 20 km of Kings ton Harbor, an active port for hundreds of years (Long 1774), and the tradition of active horticulture on this island (Adams 1972) suggest the possibility of recent introduction.Many other natural disjunct patterns of distribution of closely related taxa between Jamaica and tropical America are well documented (Howard 1973). The morphologically distinct (albeit weak) nature of S. foetida var. jamaicensis suggests this as a distinct and indigenous variety.

18c. Simsia foetida var. megacephala (Schultz-Bipontinus ex S. F. Blake) D. M. Spooner, comb. et stat. nov. Simsia megacephala Schultz-Bipontinus ex S. F. Blake, Proc. Amer. Acad. Arts 49: 391. 1913.-TYPE: MEXICO:culti vated plant grown from achenes of Liebmann 546 in the Botanic Garden of Deidesheim, Rhine-Palatinate, 25Nov 1859, Schultz-Bipontinus s.n. (holo type:GH!, photo: ENCB!; isotypes: BM! P[2]!). Simsia cronquistii H. Robinson & R. D. Brettell, Phytologia 24: 370. 1972. TYPE: MEXICO. Oaxaca: 97 mi SE of Oaxaca and 54 mi NW of Tehuantepec, 3100 ft, along the Pan-American Hwy in small-forest region, 13 Oct 1962, Cronquist 9661* (holotype: US!; isotypes: GH! MO! MICH! MSC! NY! SMU! TEX!).

Stems 1-3 m tall. Leaves with petioles 1-5.5 cm long, some leaves, especially the upper, often winged throughout with wings often 1 cm wide or more, at base often cordate-clasping, sometimes united from those of opposite leaves to form nodal discs, blades 3-12.5 cm long, 2-9 cm wide, densely glandular-puberulent with stipitate glands, setose, sometimes hispid. Peduncles 1-14 cm long. Heads 10 16 mm long, 25-35 mm in diameter. Involucre 8-13 mm long, 8-15 mm in diame 70 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 30

ter; phyllaries 19-29, 3-seriate, hirsute and glandular-puberulent; outer phyllaries 10-20 mm long, 1.6-3.2 mm wide; inner phyllaries 11.6-20 mm long, 1.4-2.7 mm wide. Pales 9.2-14 mm long, 1.9-3.1 mm wide. Ray florets 7-14; ligules 5.3-10 mm long, 2.3-4.8 mm wide, light lemon-yellow; ovary 2.3-4.8 mm long. Disc florets 35-55, corolla 5.6-8.7 mm long, tube 0.8-2.5 mm long; achenes 4.4-7.4 mm long, 2.6-4.1 mm wide, awns 4.4-7.4 mm long. Chromosome number: n = 17. Phenology. Flowering from June throughNovember. Distribution (Fig. 21). Confined to eastern Oaxaca, Mexico, on the lower slopes of the eastern end of the Sierra Madre del Sur and adjacent lowlands about the Isthmus of Tehuantepec; along roadsides and in disturbed fields in areas of tropical deciduous forest; 0-950 m.

ADDITIONAL SPECIMENS EXAMINED. Mexico. OAXACA: 12 mi NW of Tehuantepec, Hartman & Funk 4190* (MO, OS, TEX); Jalapa del Marquds, Herndndez M.118* (MEXU), Spooner & Dorado 2669* (BM, MEXU, OS, UC); Rt 190, 0.9 mi NW of Las Majadas, Jansen & La Duke 403* (ASU, ENCB, OS); Rt 190, 6.9 mi NE of Juchitan de Zaragoza, Keil & Luckow 15553* (OBI, OS); Rt 190, 10-15 km NW of Tehuantepec, King 768 (MICH); Sta. Cruz, Liebmann 546 (C, P); Salina Cruz, Ortega 25 (MEXU); Rt 195, 1 mi W of bridge over Rfo Tehuantepec, Spooner 2814* (MEXU, OS, UC); Rt 195, 4.8 mi SE of La Coyula, Spooner 2815* (MEXU, OS); Rt. 195, 3 mi SW of town center of La Coyula, Spooner & Dorado 2667* (BM, GH, MEXU, MICH, NY, OS, TENN, US, WIS); Rt 195, 3.3 mi SW of La Reforma, Spooner & Dorado 2668* (MEXU, OS, P, TEX); 8 mi NW of Tehuantepec, Webster & Armbruster 21031 (TEX, UC); Huazantlan del Rio, Mpio. SanMateo delMar, Zizumbo & Colunga 288 (CAS, TEX); ex Hort. Cantab., from seeds Hort. Paris, collector unknown, 1866 (GH, MO, NY).

Intermediates between Simsia foetida var. megacephala and S. foetida var. foetida occur (e.g., Funk & Ramos 257, King 768, Ortega 25). Most specimens of S. foetida var. megacephala are well distinguished by thewide, frequently hourglass shaped petioles, especially in the upper nodes, but some petioles are widened only at the base and sometimes fused to form nodal discs. In addition, the pales of this variety are slightly longer than those of all the others, often slightly extending above the disc florets. Although this variety is narrowly restricted in its range, it is a dominant and abundant weed at some sites. The type of S. foetida var. megacephala is a cultivated plant grown from achenes of Liebmann 546, collected in December 1841 at "Sta. Cruz." This is a common place name in much of Mexico and Oaxaca and matches at least two localities within the likely range of var. megacephala. The name also is a possible transcription error for "Salina Cruz," also in Oaxaca within its range. Liebmann's itineraries (Oersted 1867, 1869; Hemsley 1887) make no mention of "Sta. Cruz." Also, Liebmann was possibly in northern Oaxaca at that time, outside the known range of this variety. His later excursions in 1842, however, traversed much of the range of the variety, and the year 1841 is possibly an error for 1842.

18d. Simsia foetida var. grandiflora (Bentham) D. M. Spooner, comb. et stat. nov. Simsia grandiflora Bentham inOersted, Vidensk. Meddel. Dansk Natur hist. Foren. Kj0benhavn 1852: 92. 1853. Encelia grandiflora (Bentham) Hemsley, Biol. cent.-amer., Bot. 2:184. 1881.-TYPE: [NICARAGUA:Chinan dega]: In dry fields at foot of mont El Viejo [Volcan El Viejo, Volcan San Cristoball, 1851, Oersted 100 (holotype: K!, fragment:US!, photos: GH! OS! US!; isotype: C!). 1990 SIMSIA 71

Simsia polycephala Bentham inOersted, Vidensk. Meddel. Dansk Naturhist. Foren. Kj0benhavn 1852: 93. 1853. Encelia polycephala (Bentham) Hemsley, Biol. cent.-amer., Bot. 2: 184. 1881.-TYPE: [COSTARICA]. Guanacaste: In dry fields between Sap6a and Tortuga, 1851, Oersted 160 (holotype: K!, fragment:US!, photos: GH! OS! US!; isotypes: C[2]!).

Stems 1-4 m tall. Leaves with petioles 1-10 cm long, unwinged or winged, at base sometimes auriculate-clasping, or with stipules 0-2 cm long, these sometimes united with those of opposite leaves to form nodal discs; blades 3-14 cm long, 2-15 cm wide, moderately to densely glandular-puberulent with subsessile glands, setose, sometimes hispid. Peduncles 1-14 cm long. Heads 13-19 mm long, 25-45 mm in diameter. Involucre 9-13 mm long, 10-22 mm in diameter; phyllaries 21-66, 3-4-seriate, hirsute and glandular-puberulent; outer phyllaries 8-14 mm long, 1.9 3.5 mm wide; inner phyllaries 8.4-14.5 mm long, 1.2-3 mm wide. Pales 8-11.7 mm long, 1.8-2.8 mm wide. Ray florets 11-45, ligules 7.6-12 mm long, 3-7 mm wide, mostly orange-yellow, sometimes lemon-yellow in central and eastern Guatemala; ovary 6.1-8 mm long. Disc florets 66-172, corolla 6.1-7.4 mm long, tube 0.9-1.7 mm long; achenes 4-6.2 mm long, 2.5-3.4 mm wide, awns 2.7-6 mm long. Chromo some number: n = 17. Phenology. Flowering fromOctober throughMarch. Distribution (Fig. 21). Guatemala to Costa Rica; in dry weedy thickets and streamsides in areas of tropical deciduous forests; 0-1700 m.

REPRESENTATIVE SPECIMENS. Guatemala. CHIQUIMULA: between Jocatan and Camotan, Molina & Molina 25217* (C, F, NY, US); Esquipulas-Chiquimula Rd, 5 mi SE of bridge over river at San Jacinto, Spooner & Dorado 2740* (B, BM, GH, MEXU, MICH, NY, OS, TENN, UC).-JUTIAPA: along Pan American Hwy, 3mi W of San Jose Acatempa, Spooner & Dorado 2695* (MEXU, OS, TEX, UC, US); 2.5 mi N of Pan-American Hwy on rd to Chiquimula, Spooner & Dorado 2697* (MEXU, OS, TEX). SANTAROSA: along rd SE of Barbarena, Standley 77857 (F). El Salvador. AHUACHAPAN:vicinity of Ahuachapan, Standley 19947 (NY, US).-LA LIBERTAD:Col6n, Williams & Molina 15188* (F, GH). LA UNI6N: Gulf of Conchagua, Barclay 2600 (BM, MICH, US).-SAN SALVADOR: vicinity of Tonacatepeque, Calder6n 250 (GH, NY, US), Standley 19483* (GH,MO, US); airport at San Salvador, Heiser R7 (IND).-SAN VINCENTE:vicinity of San Vincente, Standley & Padilla 3549* (F).-SANTA ANA: vicinity of Metapan, Standley & Padilla 3179 (F). Honduras. CHOLUTECA:between El Espino & San Marcos de Col6n, Molina 23238* (G, NY); along Pan-American Hwy, between El Chinchayote and Comali, Molina & Molina 24566* (DUKE, F, NY, US).-EL VALLE: along Pan-American Hwy, 6 mi S of JicaroGalan, Spooner & Dorado 2699* (EAP, MEXU, OS, TENN, US). Nicaragua. ESTELf:4 km on way to San Juan Limay, Molina 23157* (F, MO, NY, S); Paso Le6n, 5 km from Esteli on rd to Miraflor, Moreno 22322 (MO, OS); Pan-American Hwy, 2 mi N of center of Esteli, Spooner & Dorado 2700* (MEXU, OS, TEX, UC); Pan-American Hwy, 14.6 mi N of bridge over Rio Esteli on N side of Esteli, Spooner & Dorado 2702* (MEXU, OS, TEX).-GRANADA:Niquinohomo, Baker 2419* (G, GH, MO, MSC, NY, TEX, UC, US, W); km 54, rd to Diria, Morena & Stevens 22882 (MO, OS).-LE6N: Cafi6n of Rio Sinecapa, near Santa Rosa, Williams & Molina 42449 (F,MICH).-MANAGUA: Sierra de Managua, Grant 7339 (F, GH); along Hwy 2, 2 km SSW of Hwy 12, Stevens 4768 (MO, OS). MATAGALPA:Pan-American Hwy, 0.9 mi S of rd to Jinotega, Spooner & Dorado 2703* (C,MEXU, OS, OSH, US); Pan-American Hwy, 7.0 mi S of bridge over Rio Grande at Sebaco, Spooner & Dorado 2723* (MEXU, OS, US); 27.6 mi S of Sebaco, Stuessy & Gardner 4552 (CHAPA, ENCB, MO, OS, WIS). Costa Rica. CARTAGO: Ojos de Agua, ex herb. Klatt 352 & 382 (GH).-GUANACASTE: Rio Guapote, Santa Rosa National Park, Janzen 10652* (MO); between Capalchi and La Cruz, 29 Jan 1909, Brenes s.n. (NY).-PUNTARENAS:Isla de Chira, Valerio 1507* (F).

Simsia foetida var. grandiflora has the largest heads of any species within the genus. This variety and S. foetida var. panamensis also differ from other varieties of 72 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 30

S. foetida by the prevalence of orange-yellow ligules and phyllaries that tend to be more indurate below. Intermediates between var. grandiflora and var. foetida occur in central and eastern Guatemala.

18e. Simsia foetida var. panamensis (H. Robinson & R. D. Brettell) D. M. Spooner, comb. et stat. nov. Simsia panamensis H. Robinson & R. D. Brettell, Phytologia 24: 372. 1972.-TYPE: PANAMA.Panama': Bella Vista, "Silvu laca,"brushy field, 28Nov 1923, Standley 25386 (holotype:US!, photo: OS!)

Stems 1-4 m tall. Leaves with petioles 1-8 cm long, without stipules or with stipules at the base 0-1.5 cm long, these sometimes united with those of opposite leaves to form nodal discs; blades 3-18 cm long, 2-14 cm wide, pilose to subsericeous and glandular-puberulent with subsessile glands. Peduncles 1-13 cm long.Heads 12-16 mm long, 25-35 mm in diameter. Involucre 9-13 mm long, 9-18 mm in diameter; phyllaries 30-44, 3-seriate, pilose and glandular-puberulent; outer phyllaries 6-10 mm long, 1.9-3.5 mm wide; inner phyllaries 7.2-10.5 mm long, 1 2.8 mm wide. Pales 6.5-8.9 mm long, 1.3-2.8 mm wide. Ray florets 12-25; ligules 5.1-11 mm long, 2.2-5.2 mm wide, predominantly orange-yellow, but varying to light lemon-yellow; ovary 3.5-6.4 mm long. Disc florets 40-151, corolla 5.2-6.9 mm long, tube 0.7-1.5 mm long; achenes 3.1-5.1 mm long, 1.7-2.5 mm wide; awns 2.2-5 mm long. Chromosome number: n = 17. Phenology. Flowering fromNovember throughApril. Distribution (Figs. 14, 21). Panama City, Panama, southwest to the Azuero Peninsula, with disjunct populations inVenezuela and Bolivia; roadsides and old fields in areas of tropical deciduous forests; 0-500 m.

REPRESENTATIVE SPECIMENS. Panama. COCLt: entrance to Rio Hato airbase, Blum & Tyson 1866* (MO); Rio Hato Airstrip, Dwyer 1146* (MO, GH, UC, US); Aguadulce, Pittier 4847* (US); 6.5 km SW of Penonome, Spooner 2904* (OS, PMA, QCA); 4 mi W of Ant6n, Tyson & Blum 2595* (MO). HERRERA: outskirts of Pese on rd to La Avena, Burch et al. 1312* (F, MO, UC, US); 4 mi S of Los Pozos, Dwyer 2697* (MO); along dirt rd near Pan-American Hwy, 2.5 mi W of Divisa, Spooner 2905* (OS, QCA).-Los SANTOS: 7 mi S of Chitrd, Croat 9703* (MO); 17.8 mi S of Macaracas, Lewis et al. 1616* (GH, MO, US); Monagre Beach, 5 mi SE of Chitre, Tyson et al. 3024* (MO); 12 mi S of Macaracas, Tyson et al. 3069* (MO).-PANAMA: 1 mi W of Rio Cot6n, D'Arcy 9756* (MO); El Valle, Harvey 5166* (F); Sabanas, Paul 595* (LL, US); Punta Paitilla, Piper 5427* (NY, US); Pan-American Hwy, 2 km NE of turnoff to La Chorrera, Spooner 2902* (OS, PMA, QCA, UC); 1 km SW of San Carlos, Spooner 2903* (OS, PMA, QCA, US).-VERAGUAS: Pan-American Hwy, 10.9 km W of jct with rd to Chupampa, near border with Herrera State, Spooner 2906* (OS, QCA). Venezuela. GUARICO:Guardatinajas, Crespo 1983* (MY, OS). Bolivia. SANTACRUZ: between Charagua & Izozog, Cdrdenas 2666 (US). TARIJA:Fortin Crevaux, Fries 1645 (S).

Simsia foetida var. panamensis shares many traits with S. foetida var. grandi flora. It is generally easily identified by its pilose leaves and smaller achenes. An intermediate with S. foetida var. grandiflora from Costa Rica, Brenes s.n., listed under S. foetida var. grandiflora, combines characters of these two varieties. The Venezuelan collection appears to be an isolated disjunct, but it is from a large population and is not a waif. Because it occurs in an agricultural area, I suggest it may have been recently introduced. The two collections from Bolivia are very similar to other representatives of this variety but have more gray-green leaves, dark-brown to black anthers, and achenes up to 7 mm long. Although these Boliv 1990 SIMSIA 73 ian populations may represent a new taxon in Simsia, they are not so treated here because of their similarity to S. foetida var. panamensis and because littlematerial exists for adequate comparisons.

DOUBTFUL AND EXCLUDED NAMES

Simsia sect. Geraea (Torrey & A. Gray ex A. Gray) A. Gray, Mem. Amer. Acad. Arts 1 (PI. fendl.): 85. 1849.-TYPE: Simsia canescens (Torrey& A. Gray ex A. Gray) A. Gray. = Geraea Torrey & A. Gray ex A. Gray, fide Blake (1913).

Simsia abrotanoides (Meisner) F. v. Mueller, Syst. census Austr. pl. 66. 1882. abrotanoidesMeisner, P1. Preiss. 1: 517. 1845.-SYNTYPES:ad ripam flum. Cygnorum, 30 Sep 1839, Preiss 2622 (not located); Swan River, Drummond 587 (not located). = Stirlingia abrotanoidesMeisner, fide Jackson (1895).

Simsia anethifolia R. Brown, Trans. Linn. Soc. London 10: 153. 1810.-TYPE: .Victoria: southern coast, "Lewins Land" [on the southern main land adjacent to Bass Strait, fide Brown, 1810], [1802, fide Stearn, 1960], Brown s.n. (not located). = Stirlingia tenuifolia (R. Brown) Steudel, fide Jack son (1895).

Simsia canescens (Torrey & A. Gray ex A. Gray) A. Gray, Mem. Amer. Acad. Arts 1 (P1. fendl.): 85. 1849.-TYPE: U.S.A. :interior of California, 1844, Fremont 393 (lectotype, here designated: GH). = Tor rey & A. Gray ex A. Gray, fide Blake (1913).

Simsia frutescens A. Gray in Torrey, Rep. U.S. Mex. bound. (House ed.) 2: 89. 1859 [1858, see Stafleu and Cowan, 1976, entry no. 1670 under Emory, W. H.].-TYPE: U.S.A. CALIFORNIA:Sierra Prieta, near Fort Yuma, Dec 1854, Schot s.n. (lectotype, here designated: GH). = (A. Gray) A. Gray, fide Blake (1913).

Simsia heterophylla (Cavanilles) Persoon, Syn. pl. 2: 478. 1807, non Simsia heterophylla (Humboldt, Bonpland & Kunth) de Candolle, 1836. Coreopsis heterophylla Cavanilles, Icon. 3: 34 t. 268. 1795. Ximenesia cavanillesii Sprengel, Syst. veg. 16(3) 605. 1826. Echinacea heterophylla (Cavanilles) D. Don in Sweet, Brit. fl. gard. ser. 2. 1: pl. 32. 1831 [18301.-TYPE:Cultivated in the Royal Botanic Garden of Madrid, from unvouchered achenes from the Sesse & Mocinlo expedition; Cavanilles's plate fixes the application of the name. = Iostephane heterophylla (Cavanilles) Bentham ex Hemsley, fide Strother (1983).

Simsia hispida (Humboldt, Bonpland & Kunth) Cassini, Dict. Sci. Nat. 59: 137. 1829.Ximenesia hispida Humboldt, Bonpland & Kunth, Nov. gen. sp. 4 [folio]: 178. 1818; 4 [quarto]: 227. 1820. Encelia hispida (Humboldt, Bonpland & Kunth) Hemsley, Biol. cent.-amer., Bot. 2: 183. 1881.-TYPE: MEXICO. Micho acan: Valladolid [Morelia, 6200 ft., Sep 1803, fide Sprague, 19241,Bonpland 74 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 30

4277 (holotype: P-HBK, microfiche: IDC 6209. 109. I. 1!). = Viguiera sessili folia de Candolle, fideBlake (1918b).

Simsia latifoliaR. Brown, Suppl. prod. fl.Nov. Holl. 9. 1830.-TYPE: AUSTRALIA. : King Georges's Sound, 1823, Baxter s.n. (not located). = (R. Brown) Steudel, fide Jackson (1895).

Simsia scaposa A. Gray, Smithsonian Contr. Knowl. 5, Art. 6 (P1.wright. 2): 88. 1853.-TYPE: U.S.A. New Mexico: between theMimbras and theRio Grande, Oct 1851, Wright s.n. (holotype: GH). = Encelia scaposa (A. Gray) A. Gray, fideBlake (1913).

Simsia simplex (Lindley) F. v.Mueller, Syst. census Austral. pl. 66. 1882. Stirlingia simplex Lindley, Sketch veg. Swan R. 30. 1839.-TYPE: not located. = Stir lingia simplex Lindley, fide Jackson (1895).

Simsia tenuifoliaR. Brown, Trans. Linn. Soc. London 10: 152. 1810.-TYPE: Aus TRALIA.Victoria: southern coast, "Lewins Land" [on the southern mainland adjacent to Bass Strait, fideBrown, 1810] [1802, fide Stearn, 1960], Brown s.n. (not located). = Stirlingia tenuifolia (R. Brown) Steudel, fide Jackson (1895).

Simsia teretifolia (Meisner) F. v. Mueller, Syst. census Austral. pl. 66. 1882. Stirl ingia teretifoliaMeisner, P1. Preiss. 1: 515. 1845.-TYPE: in regionibus inte rioribusAustraliae meridionali-occidentalis, Feb 1841, Preiss 768 (not located). = Stirlingia teretifoliaMeisner, fide Jackson (1895).

ACKNOWLEDGMENTS

I gratefully acknowledge numerous people for their help throughout this work. Tod F. Stuessy, my major professor, suggested the problem and has consistently provided help and encouragement through out all phases of this work and my graduate study. I also extend appreciation to my committee members, Daniel J. Crawford, Gary L. Floyd, and Thomas N. Taylor, for their advice and critical reading of the manuscript. Numerous friends and colleagues provided help in various ways. David M. Brandenburg, Edward E. Schilling, and Charles M. Triplehorn all provided funds and one month each of their time in the field. Armando Burgos, Oscar R. Dorado R., and Jose L. Villasefior R. accompanied me in Mexico, Jose Guevana B. in Colombia, and Andrea Clausen in Argentina. Wayne J. Elisens, Katherine L. Gross, Paul 0. Lewis, Donald H. Les, John J. Furlow, Ho-Yih Liu, and Richard Whitkus provided statistical advice. Theodore Barkley and Harold Robinson provided valuable discussions on species limits and generic problems in Simsia. Wayne J. Elisens and Stephen J. Wolf provided photographic assistance. Elton F. Paddock helped in the interpretation of meiotic configurations. Dale Johnson helped locate obscure literature references. Michael 0. Dillon provided cytological material of Simsia dombeyana from Peru. Christiane Anderson, John H. Beaman, Edward E. Schilling and John L. Strother provided invaluable help in reviewing the manuscript. Others aided in numerous ways, including valuable discus sions, the loan of specimens, help with obtaining collecting permits, or welcoming me into their homes while I was in foreign countries. They include Mireya D. Correa A., Arthur Cronquist, Santiago Diaz P., Luis D. G6mez P., Robert K. Jansen, David J. Keil, Stephen D. Koch, Thomas G. Lammers, John F. Pruski, T. P. Ramamoorthy, Mario Sousa S., Billie L. Turner, and Thomas L. Wendt. David Dennis of the College of Biological Sciences made the line drawings. The Ohio State University Instructional and Research Computer Center provided computer time. 1990 SIMSIA 75

Grant support was provided by a Doctoral Dissertation Improvement Grant from the National Science Foundation (BSR-8313268) to The Ohio State University (Dr. Tod F. Stuessy, principal investi gator), The Society of Sigma Xi, The Ohio State University Graduate School (Alumni Research Award), The Ohio State University Department of Botany, and theAmerican Society of Plant Taxono mists (Herbarium Travel Award). Appreciation is expressed to The Ohio State University for the award of a Presidential Fellowship, which provided me with an unencumbered final year to complete thiswork. I thank the curators of the following herbaria for the use of specimens:ARIZ, ASU, B, BAB, BM, BR, C, CAS, CGE, CHAPA, CM, COL, CR, DES, DS, DUKE, ENCB, F, Fl, G, GH, GOET, IJ, IND, JE, K, L, LE, LIL, LL, LPB, M, MA, MASS, MEXU, MICH, MPU, MSC, MY, NMC, OBI, OS, OSH, OXF, P, PH, PMA, POM, Q, QCA, RSA, S, SI, SMU, SRSC, TENN, TEX, U, UC, UPS, US, VEN, and W. Finally, I thankmy wife, Ann K. Spooner, without whose patience, support, and care of our thiswork never would have taken place.

LITERATURE CITED

Adams, C. D. 1972. Flowering Plants of Jamaica.Mona: University of theWest Indies. Barkley, T. M. 1988. Variation among the aureoid Senecios of North America: A geohistorical interpre tation. Bot. Rev. 54: 82-105. Bentham, G. 1853. Compositae centroamericanae: Bestemmelser og Beskrivelser af G. Bentham. Vidensk. Meddel. Dansk Naturhist. Foren. Kj0benhavn. 1852: 65-114. Blake, S. F. 1913. A revision of Encelia and some related genera. Proc. Amer. Acad. Arts 49: 346-396. 1917. New and noteworthy Compositae, chieflyMexican. Contr. Gray Herb. 52: 16-59. 1918a. New spermatophytes collected inVenezuela and Curacao by Mssrs. Curran & Haman. Contr. Gray Herb. 53: 30-55. 1918b. A revision of the genus Viguiera. Contr. Gray Herb. 54: 1-205. 1924. New American . Contr. U.S. Natl. Herb. 22: 587-661. 1926. Bertoloni's Guatemalan Asteraceae. Bull. Torrrey Bot. Club 53: 215-218. 1928. Twelve new American Asteraceae. J.Wash. Acad. Sci. 18: 25-37. Brauner, S., and L. D. Gottlieb. 1987. A self-compatible plant of Stephanomeria exigua subsp. co ronaria (Asteraceae) and its relevance to the origin of its self-pollinating derivative S. malheurensis. Syst. Bot. 12: 299-304. Brown, R. 1810. Prodromusflorae Novae Hollandiae. London: Johnson and Co. Burdet, H. M. 1973. Cartulae ad botanicorum graphicem. III. Candollea 28: 407-440. Burkart, A. 1974. Flora ilustrada de Entre Rios (Argentina), Tomo VI, Parte VI: Dicotiledoneas Metaclamideas (Gametopetalas), B: Rubiales, Cucurbitales, Campanulales (incluso Compuestas). Buenos Aires: Colecci6n cientifica del I.N.T.A. Burtt, B. L. 1970. Intraspecific categories in flowering plants. Biol. J. Linn. Soc. 2: 233-238. Cabrera, A. L. 1978. Flora de la provincia de Jujuy, Republica Argentina, Tomo XIII, Parte X: Compositae. Buenos Aires: Colecci6n cientifica del I.N.T.A. Candolle, A. P. de 1836. Compositae. In Prodromus systematis naturalis regni vegetabilis, ed. A. P. de Candolle, 5: 4-695. Paris: Treuttel etWurtz. Cantino, P. D. 1982. A monograph of the genus Physostegia (Labiatae). Contr. Gray Herb. 211: 1-105. Cassini, H. 1829. Simsia. In Dictionnaire des sciences naturelles, ed. F. Cuvier, 59: 135-137. Paris: Le Normant. Cavanilles, A. J. 1791. Icones et descriptiones plantarum 1(3): 53-67. Madrid. Clark, C., and D. L. Sanders. 1986. Floral ultraviolet in the Encelia alliance. Madronio 33: 130-135. Crawford, D. J., and T. F. Stuessy. 1981. The taxonomic significance of anthoclors in the subtribe Coreopsidinae (Compositae, Heliantheae). Amer. J. Bot. 68: 107-117. Cronquist, A. 1978. Once again, what is a species? In Beltsville Symposium inAgricultural Research Biosystematics in Agriculture, eds. J. A. Romberger et al., 3-20. Monclair: Allanheld and Osmun. . 1981. An integratedsystem of classification of flowering plants. New York: Columbia University Press. . 1985. History of generic concepts in the Compositae. Taxon 34: 6-10. Cuatrecasas, J. 1954. Notas a la flora de Colombia, XIII. Revista Acad. Colomb. Ci. Exact. 9: 233-249. 76 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 30

D'Arcy, W. G. 1975. Sigesbeckia, Simsia. In Flora of Panama, ed.W. G. D'Arcy. Ann. Missouri Bot. Gard. 62: 1132-1138. Dillon, M. 0. 1984. A systematic study of Flourensia (Asteraceae, Heliantheae). Fieldiana, Bot. n.s. 16: 1-66. Dixon, W. J., M. B. Brown, L. Engelman, J. W. Frane, M. A. Hill, R. I. Jennrich, and J. D. Toporek. 1985. BMDP statistical software. Berkeley: University of California Press. Dugand, A. 1944. Itinerarios botanicos de Jose Jer6nimo Triana. Revista Acad. Colomb. Ci. Exact. 5: 483-489. Fahn, A. 1982. Plant Anatomy. 3d ed. Oxford: Pergamon Press. Fernald, M. L. 1897. Some rare and undescribed plants collected by Dr. Edward Palmer, atAcapulco, Mexico. Proc. Amer. Acad. Arts 12: 86-94. Funk, V. A. 1982. The systematics of Montanoa (Asteraceae, Heliantheae). Mem. New York Bot. Gard. 36: 1-133. 1985. Phylogenetic patterns and hybridization. Ann. Missouri Bot. Gard. 72: 681-715. Grant, V. 1954. Genetic and taxonomic studies inGilia. IV. Gilia achilleaefolia. Aliso 3: 1-18. . 1981. Plant Speciation. 2d ed. New York: Columbia University Press. Gray, A. 1846. The characters of some new genera and species of Compositae from Texas. Proc. Amer. Acad. Arts 1: 46-50. 1849. Plantae fendlerianae novi-mexicanae. Mem. Amer. Acad. Arts 1: 1-116. 1850. Plantae lindheimerinae. Boston J. Nat. Hist. 6: 210-264. 1853. Plantae wrightianae texano-neo-mexicanae, 1. Smithsonian Contr. Knowl. 5 (Art. 6): 1-119. . 1873. Notes on Compositae and characters of certain genera and species, etc. Proc. Amer. Acad. Arts 8: 631-661. . 1883. Characters of new Compositae, with revisions of certain genera, and critical notes. Proc. Amer. Acad. 19: 1-146. . 1887. In S. Watson, List of plants collected by Dr. Edward Palmer in the state of Jalisco, Mexico, in 1886. Proc. Amer. Acad. Arts 22: 396-481. Greenman, J. M. 1903. New and otherwise noteworthy angiosperms from Mexico and Central America. Proc. Amer. Acad. Arts 39: 69-120. Harrington, H. D., and L. W. Durrell. 1957. How to identifyplants. Chicago: Swallow Press. Heiser, C. B., Jr., and D. M. Smith. 1955. New chromosome numbers in Helianthus and related genera (Compositae). Proc. IndianaAcad. Sci. 64: 250-253. Heiser, C. B., Jr., D. M. Smith, S. B. Clevenger, and W. C. Martin, Jr. 1969. The North American sunflowers (Helianthus).Mem. Torrey Bot. Club 22: 1-218. Hemsley, W. B. 1881. Encelia. InBiologia centrali-americana.Vol. 2, ed. E. D. Godman and 0. Salvin, 183-185. London: Porter and Dulan and Co. . 1887. A sketch of the history of the botanical exploration of Mexico and Central America. Biologia centrali-americana. Vol. 4, ed. E. D. Godman and 0. Salvin, 117-137. London: Porter and Dulan and Co. Hess, R. 1938. Vergleichende Untersuchungen uber die Zwillingshaare der Compositen. Bot. Jahrb. Syst. 68: 435-496. Hieronymus, G. 1901. Compositae. Bot. Jahrb. Syst. 29: 1-85. Howard, R. A. 1973. The vegetation of theAntilles. In Vegetation and vegetational history of northern Latin America, ed. A. Graham, 1-38. Amsterdam: Elsevier Scientific Publishing Co. . 1979. External morphology, Part II. In Anatomy of the Dicotyledons. 2d ed. Vol. 1, ed. C. R. Metcalfe and L. Chalk, 16-24. Oxford: Oxford University Press. Hutchinson, J. 1973. The families of flowering plants. 3d ed. London: Oxford University Press. Jackson, B. D. 1895. Index Kewensis. Oxford. Jansen, R. K. 1985. The systematics of Acmella (Asteraceae-Heliantheae). Syst. Bot. Monogr. 8: 1-115. Jansen, R. K., and T. F. Stuessy. 1980. Chromosome counts of Compositae from Latin America. Amer. J. Bot. 67: 585-594. Kapadia, Z. J. 1964. Varietas and subspecies: A suggestion towards greater uniformity. Taxon 12: 257-259. Keil, D. J., and T. F. Stuessy. 1977. Chromosome counts of Compositae from Mexico and the United States. Amer. J. Bot. 64: 791-798. Kunth, C. S. 1818. Ximenesia. InNova genera et species plantarum by A. Humboldt, A. Bonpland, and C. S. Kunth. Folio ed., 4: 178-179. 1990 SIMSIA 77

Kuntze, C. E. 0. 1898. Encelia. Revisio generum plantarum 3(2): 145. London: Arthur Felix. Kyhos, D. W. 1967. Natural hybridization between Encelia and Geraea (Compositae) and some related experimental investigations.Madrofio 19: 33-43. La Duke, J. C. 1982. Revision of Tithonia. Rhodora 84: 453-522. Lenz, L. W. 1986.Marcus E. Jones, western geologist, mining engineer and botanist. Claremont, Califor nia: Rancho Santa Ana Botanic Garden. Levin, D. A. 1967. Variation in Phlox divaricata. Evolution 21: 92-108. Loesener, T. 1923. Plantae Selerianae. Verh. Bot. Vereins Prov. Brandenburg, 84-122. Long, E. 1774. History of Jamaica. London: Lowdnes. Maddison, W. P., M. J.Donoghue, andD. R. Maddison. 1984.Outgroup analysis and parsimony. Syst. Zool. 33: 83-103. McVaugh, R. 1956. Edward Palmer: Plant explorer of theAmerican West. Norman: University of Oklahoma Press. . 1977. Botanical results of the Sesse andMocifio expedition (1787-1803). I. Summary of excur sions and travels. Contr. Univ. Michigan Herb. 11: 97-195. . 1984. Flora Novo-Galiciana, Vol. 12. Compositae. Ann Arbor: The University of Michigan Press. Millspaugh, C. F. 1904. In Plantae Yucatanae, ed. C. F. Millspaugh andA. Chase. Publ. Field Colum bianMus., Bot. Ser. 3: 85-154. Mitsakos, A., and P. Proksch. 1985. Comparative benzofuran chemistry of Enceliopsis. Biochem. Syst. Ecol. 13: 257-259. Nash, D. L. 1976. Simsia. In Flora of Guatemala, ed. D. L. Nash and L. 0. Williams. Fieldiana, Bot. 24(12): 312-317. Nordenstam, B. 1977. Senecioneae and Liabeae-systematic review. In The biology and chemistryof the Compositae, ed. V. H. Heywood et al., 799-830. London: Academic Press. Oersted, A. A. 1867. Bidrag til egeslaegtens systematik. Vidensk Meddel. Dansk Naturhist. Foren. Kj0benhavn 1866: 11-88. . 1869. [Itinerary]. In Chenes de l'Amerique tropicaleby F. M. Liebmann, vii-x. Leipzig: Leopold Voss. Persoon, C. H. 1807. Synopsis plantarum. Vol. 2. Paris: Treuttel etWurtz. Ralston, B., G. Nesom, and B. L. Turner. 1989. Documented plant chromosome numbers for 1989: Chromosome numbers inMexican Asteraceae with special reference to the tribeTageteae. Sida 13: 359-368. Randolph, L. F., I. S. Nelson, and R. L. Plaisted. 1967.Negative evidence of introgression affecting the stability of Iris species. Cornell Univ. Agr. Exp. StationMem. 398: 1-56. Raven, P. H. 1976. Systematics and plant population biology. Syst. Bot. 1: 284-316. Rieseberg, L. H., R. Carter, and S. Zona. In press. Molecular tests of the hypothesized origin of two diploid Helianthus species. Evolution. Rieseberg, L. H., and E. E. Schilling. 1985. Floral flavonoids and ultraviolet patterns in Viguiera (Compositae). Amer. J. Bot. 72: 999-1004. Rieseberg, L. H., D. E. Soltis, and J. D. Palmer. 1988. A molecular reexamination of introgression between and H. bolanderi (Compositae). Evolution 42: 227-238. Robinson, H. 1979. Studies in the Heliantheae (Asteraceae). XVIII. A new genus Helianthopsis. Phytologia 44: 257-269. . 1981. A revision of the tribal and subtribal limits of theHeliantheae (Asteraceae). Smithsonian Contr. Bot. 51: 1-102. . 1983. A generic revision of the tribe Liabeae (Asteraceae). Smithsonian Contr. Bot. 54: 1-69. Robinson, H., and R. D. Brettell. 1972. Studies in the Heliantheae (Asteraceae). II. A survey of the Mexican and Central American species of Simsia. Phytologia 24: 361-377. Rohlf, J., J. Kishpaugh, and D. Kirk. 1972. NT-SYS. Numerical taxonomy system of multivariate statisticalprograms. Stony Brook: State University New York. Rose, J. N. 1895. Report on a collection of plants made in the states of Sonora and Colima, Mexico, by Dr. Edward Palmer, in the years 1890 and 1891. Contr. U.S. Natl. Herb. 1: 293-366. Ruiz, H. 1940. Travels of Pav6n and Dombey in Peru and Chile (1777-1788), with an epilogue and official documents added by Agustin Jesus Barreiro. Translation by B. E. Dahlgren. Field Mus. Nat. Hist., Bot. Ser. 21: 1-372. Rzedowski, J. 1983. Vegetaci6n deMexico. 2d ed. Mexico: Editorial Limusa. 78 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 30

Sanders, D. L., and C. Clark. 1987. Comparative morphology of the capitulum of Enceliopsis (As teraceae:Heliantheae). Amer. J. Bot. 74: 1072-1086. Schilling, E. E., and C. B. Heiser. 1981. Infrageneric classification of Helianthus (Compositae). Taxon 30: 393-403. Schilling, E. E., and R. K. Jansen. 1989. Restriction fragment analysis of chloroplast DNA and the systematics of Viguiera and related genera (Asteraceae: Heliantheae) Amer. J. Bot. 76: 1769-1788. Schilling, E. E., and E. M. Schilling. 1986. Chromosome numbers in Viguiera series Dentatae (Compositae). Syst. Bot. 11: 51-55. Shaw, E. E. 1987. Charles Wright on the Boundary, 1849-1852, or Plantae Wrightianae Revisited. Westport, Connecticut: Meckler Publ. Corp. Sokal, R. R., and F. J. Rohlf. 1981. Biometry. 2d ed. New York: W. H. Freeman and Company. Sprague, T. A. 1924. Humboldt and Bonpland's Mexican itinerary. Bull. Misc. Inform. 1924: 20-27. Sprengel, K. 1826. Systema vegetabilium. Vol. 3. Gottingen. Stafleu, F. A., and R. S. Cowan. 1976. Taxonomic literature. 2d ed. Vol. 1: A-G. Utrecht: Bohn, Scheltema & Holkema. . 1986. Taxonomic literature. 2d ed. Vol. 6: Sti-Vuy. Utrecht: Bohn, Scheltema & Holkema. Stearn, W. T. 1960. An introduction to Robert Brown's 'Prodromus Florae Novae Hollandiae.' In Prodromus Florae Novae Hollandiae. Reprinted, 1960.Weinheim: J. Cramer. . 1983. Botanical Latin. 3d ed. North Pomfret, Vermont: David and Charles. Stebbins, G. L. 1974. Mosaic evolution, mosaic selection and angiosperm evolution. Bot. J. Linn. Soc. 88: 149-164. Stebbins, G. L., and L. Ferlan. 1956. Population variability, hybridization, and introgression in some species of Ophrys. Evolution 10: 32-46. Straw, R. M. 1955. Hybridization, homogamy, and sympatric speciation. Evolution 9: 441-444. Strother, J. L. 1983. Pionocarpus becomes lostephane (Compositae:Heliantheae): A synopsis.Madrofio 30: 34-38. Stuessy, T. F. 1975. Milleria. In Flora of Panama, ed. W. G. D'Arcy. Ann. Missouri Bot. Gard. 62: 1058-1061. . 1977. Heliantheae-systematic review. In The biology and chemistryof theCompositae, ed. V. H. Heywood et al., 621-671. London: Academic Press. Sundberg, S., C. P. Cowan, and B. L. Turner. 1986. Chromosome counts of Latin American Compositae. Amer. J. Bot. 73: 33-38. Swofford, D. L. 1984. Phylogenetic analysis using parsimony, version 2.2 user'smanual. IllinoisNatural History Survey. Triana, M. J. 1858. Choix de plantes de Nouvelle-Grenade. Ann. Sci. Nat. Bot. ser. 4, 9: 36-52. Turner, B. L. 1978. Taxonomy of Axiniphyllum (Asteraceae-Heliantheae). Madronlo 25: 46-52. Turner, B. L., M. Powell, and R. M. King. 1962. Chromosome numbers in the Compositae. VI. Additional Mexican and Guatemalan species. Rhodora 64: 251-271. Urban, I. 1902. Notae biographicae peregrinatorum Indiae occidentalis botanicorum. Symb. antill. 3: 14-158. Wagner, W. H., Jr. 1983. Reticulistics: The recognition of hybrids and their role in cladistics and classification. In Advances in cladistics. Vol. 2, Proceedings of the second meeting of theWilli Hennig Society, ed. N. I. Platnik and V. A. Funk, 63-79. New York: Columbia University Press. Warwick, S. I., J. F. Bain, R. Wheatcroft, and B. Thompson. 1989. Hybridization and introgression in Carduus nutans and C. acainthoides reexamined. Syst. Bot. 14: 476-494. Watrous, L., and Q. Wheeler. 1981. The out-group comparison method of character analysis. Syst. Zool. 30: 1-11.

APPENDIX 1

Chromosome numbers of species, varieties, and putative natural hybrids of Simsia and Viguiera; all 2n = 1711.Vouchers are deposited at OS unless otherwise specified.

S. amplexicaulis:MEXICO. Chiapas: Tuxtla Gutierrez, Spooner 2780. Chihuahua: Buenaventura, Spooner & Schilling 2453; Cuauhtemoc, Spooner & Schilling 2455. Coahuila: Arteaga, Spooner & Schilling 2463. Hidalgo: Pachuca, Spooner & Brandenburg2229. Jalisco:Tepatitlan, Spooner & Brandenburg 1990 SIMSIA 79

2302.Mexico: Tepetlaoxtoc, Spooner & Brandenburg2232. Oaxaca: Asunci6n Nochixtlan, Spooner & Dorado 2654. San Luis Potosi: Venado, Spooner & Brandenburg 2195. GUATEMALA.Baja Verapaz: Purulhd, Spooner & Dorado 2751. Chimaltenango: Aldea El Parcelamiento, Spooner & Dorado 2684. SanMarcos: Castalia, Spooner & Dorado 2673. S. annectens var. annectens:MEXICO. Guerrero: Puerto del Gallo, Spooner 2845. Mexico: Colorines, Spooner & Villasefior 2566. S. annectens var. grayi:MEXICO. Chiapas: Chicoasan Dam, Spooner 2788. Michoacan: Tuxpan, Spooner & Brandenburg 2279;Maravatio, Spooner & Villasefior 2562. S. calva: U.S.A. Texas: Medina Co., Castroville, Spooner & Brandenburg 2177; Val Verde Co., Amistad Recreation Area, Urbatsch 928 (LL, this count obtained by Lowell E. Urbatsch and first reported here.). S. chaseae: MEXICO.Campeche: Balonchenticul, Spooner 2802. Quintana Roo: Santa Rosa, Spooner 2811. Veracruz: Carrizal, Spooner 2881. Yucatan: Santa Elena, Spooner 2804; Yukalpeten, Spooner 2807a; Buctzotz, Spooner 2809. S. dombeyana: ECUADOR.Guayas: Daule, Spooner 2907; Duran, Spooner 2908. PERU.Cajamarca: Chilete, Dillon & Dillon 4618. S. eurylepis:MEXICO. Campeche: Campeche, Spooner 2799. Nuevo Le6n: El Cercado, Spooner 2481. San Luis Potosi: Cd. Valles, Spooner & Brandenburg 2209. Tamaulipas: Aldama, Spooner 2886. Vera Cruz: Los Bafios de Carrizal, Spooner 2882. S. foetida var. foetida: MEXICO.Chiapas: San Gregorio, Spooner 2768; La Trinitaria, Spooner 2770. JALISCO:Acatlan, Spooner & Brandenburg 2292; Guadalajara, Spooner & Brandenburg 2300. Mexico: Tejupilco de Hidalgo, Spooner & Brandenburg 2264; Otzoloapan, Spooner & Villasefior 2569. Puebla: Tehuitzingo, Spooner & Dorado 2637. GUATEMALA.Baja Verapaz: Salama, Spooner & Dorado 2747. El Progreso: Pan-American Hwy near border with Guatemala State, Spooner & Dorado 2691. Guatemala: Agua Caliente, Spooner & Dorado 2688. Putative hybrid of S. foetida var. foetida x S. lagascaeformis:MEXICO. Oaxaca: San Agustin Atenango, Spooner & Dorado 2644. Puebla: Tehuitzingo, Spooner & Dorado 2635. S. foetida var. grandiflora: HONDURAS.El Valle: Jicaro Galan, Spooner & Dorado 2699. NICARAGUA. Matagalpa: Sebaco, Spooner & Dorado 2723. S. foetida var. megacephala: MEXICO.Oaxaca: La Coyula, Spooner & Dorado 2667. S. foetida var. panamensis: PANAMA.Cocle: Penonome, Spooner 2904. Herrera: Divisa, Spooner 2905. Panama: La Chorrera, Spooner 2902. Veraguas: Pan-American Hwy near border with Herrera, Spooner 2906. S. fruticulosa: COLOMBIA.Huila: Hobo, Spooner & Guevara 2923. Magdelena: SantaMarta, Spooner 2975. Narinlo: Pasto, Spooner & Guevara 2937, 2941. Tolima: Ibague, Spooner & Guevara 2972. Valle: Loboguerrero, Spooner & Guevara 2957. ECUADOR.Imbabura: Ibarra, Spooner 2916. S. ghiesbreghtii: MEXICO.Chiapas: near Navenchauk, Spooner 2781. GUATEMALA.Baja Verapaz: near Pantin, Spooner & Linder 2753. Quezaltenango: S of Quezaltenango, Spooner & Dorado 2675. San Marcos: SanMarcos, Spooner & Dorado 2674. Solola: Panajachel, Spooner & Dorado 2683. S. holwayi: GUATEMALA.El Progreso: Morazan, Spooner & Dorado 2746. Guatemala: Agua Caliente, Spooner & Dorado 2692. S. lagascaeformis:MEXICO. Chihuahua: Jimenez, Spooner & Brandenburg 2336; Cd. Camargo, Spooner & Schilling 2458. Guerrero: Chilpancingo de los Bravos, Spooner & Burgos 2603, 2606. Jalisco: San Juan de los Lagos, Spooner & Villasefior 2489. Morelos: Cuernavaca, Spooner & Burgos 2600. Oaxaca: Oaxaca, Spooner & Triplehorn 2098; Ascunci6n Nochixtlan, Spooner & Dorado 2656. Puebla: Tepexco, Spooner & Dorado 2627; Tehuitzingo, Spooner & Dorado 2636. Putative hybrid of S. lagascaeformis x S. amplexicaulis: Jalisco: San Juan de Los Lagos, Spooner & Villasefior 2491. S. molinae: NICARAGUA.Esteli: Esteli, Spooner & Dorado 2701. Managua: El Crucero, Spooner & Dorado 2704, 2714. COSTARICA. Guanacaste: Playas del Coco, Spooner 2901. S. sanguinea:MEXICO. Guerrero: Ayotzinapa, Spooner & Burgos 2607. Jalisco:Guadalajara, Spooner & Villasefior 2495; Tizapan el Alto, Spooner & Villasefior 2537. Mexico: Temascaltepec, Spooner & Villasefior 2572; El Duranzo, Spooner & Villasefior 2567. Oaxaca: Huajuapan de Le6n, Spooner & Dorado 2641a, 2649b. GUATEMALA.Sacatepequez: Antigua, Spooner & Dorado 2684b; El Tejar, Spooner 2761. Solola: Panajachel, Spooner & Dorado 2681. S. santarosensis: COSTARICA. Guanacaste: El Gallo, Spooner 2900. S. setosa: MEXICO. Nayarit: Jesus Maria, Spooner & Villasefior 2528. 80 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 30

S. steyermarkii:GUATEMALA. Baja Verapaz: SanMiguel Chicaj, Spooner & Dorado 2748. S. tenuis:MEXICO. Michoacdn: Tepalcatepec, Spooner & Villasefior 2550. S. villasennorii: MEXICO.Chiapas: Rizo de Oro, Spooner 2789. Oaxaca: El Camar6n, Spooner 2816. V. cordifolia: U.S.A. New Mexico: Hidalgo Co., S end of Hidalgo Co. on Rt 81, Spooner & Schilling 2404.

APPENDIX 2

Results of artificial crosses of Simsia that produce F1's. For each cross, the female parent is listed first.After each cross, the number of F1's is listed. The number in brackets indicates themean [range] of pollen stainability of fertile Fl's. The vouchers are deposited at OS. The following collections were used for crosses: Simsia amplexicaulis (Spooner 2780, 2794), S. annectens var. annectens (Spooner 2845), S. annectens var. grayi (Spooner & Villasefior 2562, 2566, Spooner 2788), S. calva (Stuessy 712), S. chaseae (Spooner 2804, 2811, 2881), S. eurylepis (Spooner 2841, 2882), S. foetida var. foetida (Spooner & Dorado 2637, 2688, 2693, Spooner 2769), S. foetida var. grandiflora (Spooner & Dorado 2699), S. foetida var. megacephala (Spooner & Dorado 2667), S. lagascaeformis (Spooner & Burgos 2603, Spooner & Dorado 2656), S. lagascaeformis x S. foetida var. foetida (Spooner& Dorado 2635, 2644), S. sanguinea, pink-rayed variant (Spooner & Dorado 2657), white-rayed variant (Spooner & Dorado 2681), yellow-rayed variant (Spooner & Dorado 2649b), S. setosa (Spooner& Villasenior2528), S. tenuis (Spooner & Villasefior 2550), S. villasenorii (Spooner 2789, 2816).

Intraspecific crosses

S. annectens var. grayi (2788) x var. grayi (2562) 1 (44) S. annectens var. annectens (2845) x var. grayi (2566) 2 (92 [84-98]) S. foetida var. foetida (2688) x var. foetida (2688) 2 (99 [99-99]) S. foetida var. foetida (2688) x var. megacephala (2667) 3 (61 [51-67]) S. foetida var. grandiflora (2699) x var. foetida (2688) 4 (96 [90-99]) S. foetida var. grandiflora (2699) x var. megacephala (2667) 10 (67 [11-92]) S. sanguinea, pink-rayed variant (2657) x S. sanguinea, yellow (2649b) 8 (80 [62-92]) S. sanguinea, yellow (2649b) x S. sanguinea, pink (2657) 21 (78 [55-93]) S. sanguinea, yellow (2649b) x S. sanguinea, white (2681) 5 (67 [57-78]) S. villasenorii (2789) X S. villasenorii (2816) 12 (68 [48-98])

Interspecific crosses

S. amplexicaulis (2780) x S. lagascaeformis (2656) 3 (67 [44-99]) S. amplexicaulis (2780) x S. lagascaeformis x foetida (2635) 2 (99 [99-99]) S. foetida (2688) x S. eurylepis (2882) 1 (25) S. foetida (2677) x S. eurylepis (2882) 1 (50) S. foetida (2769) x S. eurylepis (2481) 18 (35 [4-56]) S. foetida (2769) x S. eurylepis (2882) 12 (39 [31-50]) S. foetida (2688) X S. Iagascaeformis x foetida (2644) 4 (63 [42-77]) S. foetida (2699) x S. lagascaeformis x foetida (2644) 2 (71 [68-73]) S. lagascaeformis x foetida (2644) x S. foetida (2677) 3 (95 [93-99]) S. foetida (2699) x S. setosa (2528) 1 (12) S. foetida (2688) x S. tenuis (2550) 1 (49) S. Iagascaeformis (2603) X S. lagascaeformis x foetida (2635) 1 (80) S. lagascaeformis x foetida (2635) x S. lagascaeformis (2656) 7 (97 [92-99]) S. lagascaeformis x foetida (2635) x S. eurylepis (2481) 3 (62 [58-67]) S. sanguinea (2649b) x S. foetida (2699) 1 (21) S. sanguinea (2657) X S. lagascaeformis (2656) 1 (30) S. setosa (2528) x S. chaseae (2811) 3 (1 [0-2]) S. villasenorii (2789) X S. annectens (2562) 1 (53) S. villasenorii (2789) X S. annectens (2566) 1 (36) 1990 SIMSIA 81

S. villasenorii (2816) x S. annectens (2562) 4 (50 [36-58]) S. villasenorii (2816) x S. calva (712) 4 (34 [25-41]) S. villasenorii (2816) x S. chaseae (2811) 3 (6 [4-7])

NUMERICAL LIST OF SPECIES

1. S. ghiesbreghtii 12. S. calva 2. S. santarosensis 13. S. lagascaeformis 3. S. fruticulosa 14. S. eurylepis 4. S. molinae 15. S. chaseae 5. S. holwayi 16. S. dombeyana 6. S. steyermarkii 17. S. amplexicaulis 7a. S. annectens var. annectens 18a. S. foetida var. foetida 7b. S. annectens var. grayi 18b. S. foetida var. jamaicensis 8. S. tenuis 18c. S. foetida var. megacephala 9. S. villasenorii 18d. S. foetida var. grandiflora 10. S. setosa 18e. S. foetida var. panamensis 11. S. sanguinea

INDEX TO NUMBERED COLLECTIONS EXAMINED

The numbers in parentheses refer to the corresponding names in the text and in theNumerical List of Species presented above.

Abbott, R. 401 (11). Balls, E. K. 5302 (17); 5616 (17). Acosta, M., & Dorantes 529 (17). Barclay, A. 719 (12). Acosta, S. 208 (17). Barclay, G. W. 2600 (18d). Adams, C. D. 8766 (18b). Barkley, F. A. 17M867 (11); 13225 (12); 14506 Aguilar, J. I. 93 (11); 1748 (11); 1667 (18a). (12). Albers, C. 44007 (12). Barnes et al. 6 (11). Alcorn, J. 2637 (14); 2796A (14). Barnette, B. 275 (12). Allen, E. 102 (12). Barrie et al. 650 (7b). Allen, P. H. 1131 (18e). Barroso, G. M. 35113 (16). Allen, T. J. 3 (12). Bartholomew et al. 3043 (11). Alvardo, J. 540 (17). Bartlett, H. H. 10618 (12). Alvarez et al. 7661 (12); 7755 (12). Bayona, A. 69 (17). Anderson & Laskowski 3937 (11); 4323 (18a). Beck & Saenz 724 (16). Antipovich, M. 120207 (17). Bell & Duke 17013 (13). Araiza 55 (12). Berlandier, J. L. 927 (17). Archundia, R. 55 (17). Blum & Tyson 1866 (18e). Arguelles, E. 184 (17); 271 (18a); 395 (17); 608 Boege, W. 1590 (17). (18a); 1450 (17). Bollwinkel & Wunderlin 132 (13). Aristeguieta, L. 2476 (3); 2491 (3). Bonilla, R. 33 (17). Arnold et al. 57 (12). Bonpland, A. 2350 (17); 4317 (17); 4350 (17); Arsene, G. 60 (17); 149 (17); 265 (17); 356 (17); 4360 (17). 363 (17); 1764 (17); 1805 (17); 2309 (17); Bopp, M. 295 (17). 3104 (17); 5156 (17); 9937 (17); 10497 (17). Bosse, G. 8365 (11); 8369 (11). Artamanoff, G. 193 (18d). Botteri, M. 427 (17); 428 (17); 804 (13); 805 Arvizu, J. 91 (17). (17); 808 (17). Aymard, G. 412 (1). Bourgeau, E. 155 (17); 850 (17); 959 (17); 1203 Baca, J. 114 (17). (18a); 3320 (13). Badillo, V. M. 3308 (3); 5181 (3); 6420 (3); 7161 Bray, W. L. 356 (12). (3); 7611a (18e). Breedlove, D. E. 7296 (17); 9237 (18a); 9705 Bailey, L. H. 871 (11). (1); 12254 (11); 12502 (17); 12795 (17); Baker, C. F. 2419 (18d). 14424 (17); 19972 (17); 23411 (18a); 27423 82 SYSTEMATICBOTANYMONOGRAPHS VOLUME 30

(11); 28465 (11); 29249 (17); 29291 (17); (12); 51376 (12); 51726 (12); 52462 (12); 36152 (7a); 39761 (17); 39898 (11); 40919 52488 (12); 53127 (12); 53594 (12); 53791 (17); 41161 (18a); 41302 (11); 41483 (18a); (12); 55138 (12); 58183 (12). 44455 (9); 48485 (1); 48937 (18a); 52914 Cowan et al. 5736 (18a). (9); 53743 (18a); 56258 (18a); 56339 (9); Crawford et al. 1245 (17); 1275 (11); 1305 (11). 62777 (18a). Crist6bal, C. 328 (16). Breedlove & Almeda 48030 (11); 60160 (9). Croat, T. 9699 (18e); 9703 (18e); 39207 (12); Breedlove & Davidse 54889 (11); 55461 (18a). 39215 (14). Breedlove & Raven 8288 (17); 8426 (18a); 12434 Crockett, R. 8102 (12). (17); 13283 (17). Cronquist, A. 9611 (17); 9661 (18c); 9687 (9); Breedlove & Strother 46026 (11); 46072 (11); 9708 (11); 9714 (11); 10314 (7b); 11299 46090 (18a); 46143 (11); 46303 (17); 46316 (12). (11); 46621 (11); 46972 (18a); 47040 (7b). Cronquist & Fay 10816 (18a). Breedlove & Thorne 20576 (9); 30382 (18a). Cronquist & Sousa 10403 (13); 10461 (11). Breteler, F. J. 4545 (3); 4548 (3). Crutchfield & Johnston 5206 (14); 5570 (12); Broadway, W. E. 463 (3). 5902D (7b); 5951A (18a). Brunken & Perino 473A (17). Cuatrecasas, J. 6500 (3). Buchinger 497 (7b). Cuatrecasas & Barriga 1019 (3); 1933 (3). Burbidge, R. B. 75/369 (3). Cumara, L. 2146 (3). Burch et al. 1146 (18e); 1312 (18e). Cummins, G. B. 63-373 (13); 70-SA (12); 71 Burkart, A. 10492 (16); 22323 (16); 22337 (16). 256H (18a); 71-326H (13). Bush, B. 837 (12); 1665 (12). D'Arcy, W. G. 9756 (18e); 10389 (18d). Bye, R. 4392 (17); 5696 (17). D'Arcy & Croat 4165 (18e); 4171 (18e). Bye & Weber 7854 (17). Daubenmire, R. 493 (2). Cabrera & Kiesling 24908 (16). Davidse & Davidse 9610 (11). Cabrera & Marchionni 13045 (16). Davidse et al. 5802 (3); 5805 (3). Calder6n, S. 250 (18d); 998 (11); 1209 (18d). Davies & Delevoryas 448 (14). Calvert, B. 123A (13). Davies & Turner A69 (12). Calzada, J. I. 2052 (14). Deam, C. C. 73 (17). Canby, W. 135 (12). Degener, 0. 26177 (17); 26757 (15). Cardenas, M. 2666 (18e). DeJong, D. C. D. 1412 (17); 1510 (17). Carlson, M. C. 185 (18d). Demaree, D. 63355 (12). Carvajal & Sanchez 460 (17). Devender et al. 84-546 (13). Castellanos 94 (16). Diaz, C. L. 486 (11); 975 (11); 6215 (17). Cerrate, E. 5217 (16). Diaz, J. M. 153 (17); 178 (17). Chiang et al. 8963 (13); 9211 (12); 75518 (12). Diaz et al. 10330 (17). Chase, V. H. 7494 (14). Dieterle, J. V. A. 4351 (8). Cicci6, 45 (4). Diguet, M. L. 85 (11). Clare, M. 19 (12). Dillon & Baker 866A (12). Clark, 0. M. 4695 (12). Dinelli, E. 26175 (16). Clemens & Clemens 934 (12); 1000 (12). Dodds, D. 20 (11). Clokey, I.W. 1859 (17). Dombey, J. 12 (16). Clover, E. 643 (12); 1355 (12); 1672 (12). Dorantes & Acosta 196 (11). Clute 2 (18b). Dorantes et al. 1743 (15). Coleman, J. 505 (16). Duenias, C. 23 (17). Colunga & Zizumbo 21 (10). Duge, A. 12 (17); 455 (17). Connell, L. 6506 (12). Duran, M. L. 145 (17). Conzatti, C. 131 (17); 1529 (18a); 1652 (11); Earle, F. S. 439 (12); 464 (12). 4045 (11); 4281 (18a); 4761 (13); 4997 (11). Elias, J. 125 (17); 220 (17); 237 (17); 239 (17); Conzatti & Gonzalez 403 (11); 899 (11); 1002 259 (17); 260 (17); 358 (17); 434 (17); 505 (13); 1263 (13). (17). Coon, N. 132 (17). Ellenberg, H. 1206 (16); 1303 (16). Correll, D. S. 16682 (12); 31722 (12); 32370 Ervendberg, L. C. 378 (17). (12); 44302 (12). Espinosa, F. J. 71 (18a); 711 (17). Correll & Johnston 17198 (12). Espinosa, J. 22 (17). Correll & Schweinfurth 15706 (12). Espinosa & Espinosa 60 (17); 302 (17); 453 Cory, V. 2648 (12); 2660 (12); 15736 (12); 50555 (17). 1990 SIMSIA 83

Feddema, C. 2130A (18a); 2342 (17); 2780 (11); Guerra, C. 69 (17). 2866 (7a). Guerra et al. 629 (12). Fernandez & Acosta 2155 (17). Guizar et al. 194 (18a). Ferreyra, R. 5902 (16); 6037 (16); 10482 (16); Gustafsson, R. 1471 (15). 10662 (16); 12369 (16). Haenke, T. 1899 (16). Ferreyra et al. 16281 (16). Haines, H. H. 7CA (18e). Ferris & Duncan 3607 (12). Hall, E. 339 (12). Fiedler, 0. 15330 (17). Hansen & Nee 7359 (14); 7414 (14). Fisher, G. L. 104 (12); 35335 (17); 49084 (12). Harking, F. G. 497 (17). Fleetwood, R. J. 3527 (12); 11519 (12). Harley, R. M. 21718 (16). Fletcher, H. 600 (12); 602 (12); 790 (12); 1340 Harley et al. 21987 (16). (12); 1539 (12). Harling & Andersson 13338 (16). Flores, D. 122 (17). Harmon, W. E. 4668 (18a). Flores, J. 167 (17). Harmon & Dunn 5590 (12). Flyr, D. 350 (12); 472 (12); 1558 (12). Harriman, N. A. 15863 (17). Forero, E. 886 (3). Harris, W. 6953 (18b); 6989 (18b); 8228 (18b); Franco, J. 25 (17). 10001 (18b). Fries, R. E. 1166 (16); 1645 (18e). Hartman & Funk 4169 (9); 4190 (18c). Froderstrom & Hulten 91 (18a); 433 (18a); 502 Hartweg, K. T. 145 (17). (18a). Harvey, D. R. 5166 (18e). Fryxell & Bates 2074 (13). Haught, 0. F152 (16); 155 (16); 249 (16); 1928 Funk & Geman 2321 (11). (16); 6374 (3). Funk & Hill 2301 (18a). Haught & Svenson 11563 (16). Funk & Ramos 2652 (18a). Hays, S. 447 (12). Galeotti, H. G. 2092 (11); 2478 (17). Hedrick, D. C. 263 (12). Gallegos, F. 469 (17). Heiser, C. R5 (13); R7 (18d). Garcia, E. A227 (13). Heithaus, R. 168 (2); 397 (2). Garcia, E., et al. 22 (12); 138 (17). Heller, A. A. 1860 (12). Garcia, J. 34 (17); 64 (17); 718 (12); 805 (17); Henderson, N. L. 63-501 (12); 63-699 (12); 63 1717 (18a). 999 (12). Garcia, R. 17 (12); 48 (12). Henrickson, J. 6314 (13); 7549 (13); 8026 (17); Gaumer, G. F. 910 (15); 2045 (15); 2046 (15). 13036 (13); 13695 (13); 60009A (13). Gentry, A., & Forero 15458A (3). Heriberto, Bro. 222 (18e). Gentry, H. 654 (10); 822M (10); 1036 (10); 2464 Hernandez, C. 464 (17). (10); 2751 (17); 3430 (10); 18108 (17). Hernandez, E., & Martinez X15903 (17). Gentry, H., & Arguelles 18151 (17). Hernandez, E., & Sharp X391 (18a). Gentry, H., et al. 19532 (18a). Hernandez, E., et al. X2364 (17); X2378 (17). George & Snelling 73383.6YW (17). Hernandez, G., & Rodriguez 1224 (12). Ghiesbreght, A. 305 (11); 540 (17); 658 (1). Hernindez, R. M. 118 (18c). Giovannini, F. 5 (12). Herrera & McCart 7689 (12). Goodding, L. N. 207-61 (13); 289A-61 (13); Heyde & Lux 3396 (17); 3810 (18a); 6160 (18a). 443-58 (13). High, M. M. 54 (12). Gonzalez, G. 71 (11). Hill, S. 10624 (12). Gonzalez, J. 614 (3). Hinkley, L. C. 457 (12); 3038 (12); 3966 (12). Gonzalez, L. 1389 (17); 3180 (17). Hinton, G. B. 1741 (8a); 1874 (11); 2216 (18a); Gonzalez et al. 215 (3). 2617 (18a); 2693 (7a); 2801 (7a); 2851 (11); Gould, F. 5691 (12); 5731 (12); 6971 (12). 2935 (7a); 5538 (18a); 5549 (13); 6706 (11); Graham & Johnston 4098 (14); 4281P (12); 4380 6948 (18a); 8578 (13); 8582 (18a); 9625 (12). (18a); 9818 (11); 10647 (11); 11271 (7a); Grant, V. 7339 (18d). 11651 (7a); 12714 (7a); 12885 (7a); 13260 Grashoff, J. 154 (13); 259 (17). (7b); 13289 (18a); 13320 (11); 13487 (11); Greenman, J. M. 33 (17). 14843 (7a); 14976 (7a); 15494 (17); 17475 Greenman & Greenman 5109 (18e); 5119 (18e). (12); 17681 (12). Gregg, J. 20 (17); 47 (12); 85 (12); 555 (17); 649 Holway, E. W. D. 624 (18a); 854 (5); 3740 (13); (17). 3747 (13); 5092 (18a); 5340 (13); 5360 Grinstein, S. 142 (17). (18a). Groth, B. 143 (12). Huitr6n, G. 7 (13). 84 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 30

Humbert et al. 27111 (3). Kral, R. 24777 (12). Iltis, H. H. 27054 (12); 27093 (12). Krapovickas & Schinini 35947 (16). Iltis& Cochrane 228 (17). Kruse, H. 1225 (11). Iltis& Koeppen 1257 (11). La Duke et al. 375 (13). Iltis et al. 6 (12); 544 (17); 710 (7b); 1189 (11). Lahitte, R. 49586 (16). Jackson, J. 122 (12). Lane & Longstreth 2734 (17). Jackson, R. C. 6705 (12). Lane & Olsen 2098 (11); 2104 (13). Jahn, A. 651 (3); 677 (3); 682 (3). Lanfranchi, A. 1304 (16); 1314 (16). Jansen & La Duke 390 (17); 403 (18c). Langman, I. K. 3143 (13); 3164 (13); 3552 (17); Janszen 61 (12). 3675 (18a). Janzen, D. 10652 (18d). Lape, F. 9A (17). Jermy, C. 51 (12); 863 (12). Latorre, D. L. 51 (12); 62 (12); 131 (12). Jimenez, F. 34 (17); 208 (17). Laughlin, R. M. 111 (17); 251 (18a); 988 (17); Jimenez, J. 410 (12). 1442 (17); 1711 (17); 2191 (17); 2300 (17); Johnson, J. 1068 (12); 1533 (12). 2386 (17); 2476 (17); 2875 (18a); 2959 (11). Johnson, J., & Barkley 16059M (12). Lehmann, F. C. 341 (3); 2964 (3); 4710 (3). Johnston, I.M. 7030 (12); 7080 (12); 7888 (13); Lehto et al. 5203 (12); 5233 (12); 5799 (12). 8559 (12); 9158 (12); 9218 (12). Le Sueur, H. 474 (12). Johnston, I.M., & Muller 431 (12); 791 (13). Letterman, G. 5 (12). Johnston, J. R. 245 (17); 250 (11); 259 (17); 362 Lewis, H. H., et al. 1616 (18e). (11); 484 (11); 491 (17); 987 (13). Lewis, S. 348 (12). Johnston, L. 41 (12). Liebmann, F. M. 492 (11); 546 (18c); 559 (13); Johnston, M. C. 53.280.51 (12); 3290 (12); 3804 560 (7b); 561 (7b); 586 (17); 587 (17); 589 (12); 4892B (12); 4933 (14); 5444 (12); 5454 (17); 601 (17). (12); 5758 (12); 5776 (14); 54481 (12); Liesner et al. 2661 (2). 541407 (12); 542349 (12). Lillo, M. 25375 (16); 71726 (16). Johnston, M. C., and Johnston 7252 (11). Lindheimer, F. J. 6 (12); 432 (12); 433 (12); 490 Johnston, M. C., et al. 3567 (12). (12); 900 (12). Jones, F. B. 51 (12). Long & Burch 3195 (14); 3225 (14). Jones, M. E. 26545 (12); 26551 (12); 27714 (17); L6pez, S. 2125 (3). 27778 (11). L6pez, W. 639 (17). Jorgensen, P. 1286 (16). L6pez et al. 10785 (17). Juzepczuk, S. 884 (18a). Lucas, M. M. 417 (12). Karwinsky, W. F. 1301 (13); 1303 (13). Lundell & Lundell 10701 (12). Keil & Canne 8986 (18a); 9081 (7b); 9128 (11); Lyonnet, E. 845 (18a). 9231 (14). MacBryde, J. F. 2604 (18e). Keil & Luckow 15331 (13); 15419 (17); 15480 MacDaniels, L. H. 501 (17); 572 (17). (13); 15491 (11); 15521 (11); 15553 (18c). MacKenzie, K. 75 (12). Keil & McGill 7994 (13). Magnon & Rodriguez 31 (12). Keil et al. 13442 (13). Mahler, W. F. 2995 (12); 3007 (12); 3410 (12); Kellerman, W. A. 4982 (1); 7497 (18a). 3435 (12); 3507 (12); 5277 (12); 5284 (12). Kelly, I. 672 (17); 793 (17). Mairet 108 (17). Kenoyer, L. A. 6 (12); 594 (13); 2371 (17). Marquez, W. 21 (17). Killip & Smith 16454 (3). Marsh, E. G. 192 (12); 1104 (12); 1759 (12); Killip et al. 38378 (3). 2248 (12); 2289 (12). Kim & Miller 1240 (12). Martinez, M. A. 1103 (17); 1143 (17). Kim et al. 970 (12). Martinez et al. 4830 (11). King, R. M. 768 (18c); 2239 (17); 2299 (13); Matuda, E. S116 (17); 19317 (17); 19357 (17); 2312 (13); 2319 (13); 2518 (13); 2918 (13); 19728 (17); 19752 (17); 21501 (17); 26422 3415 (17); 7337 (6). (17); 26927 (17); 26929 (17); 26954 (7a); King & Bishop 8586 (16); 8704 (16). 29349 (18a); 29412 (11); 29552 (17); 31621 King & Garvey 7008 (16). (11); 31733 (17); 38254 (11). King & Guevara 5817 (3); 5872 (3). Maxon & Hay 3751 (1). Kishler, J. 54 (17); 109 (17); 418 (17); 816 (17). McBryde, W. 393 (12). Klatt, A. 2964 (3). McCart, W. 237 (12). Knobloch, I.W. 5606 (17); 5990 (17); 5999 (17). McCart et al. 8 (12). Koch, R. 2750 (12). McDaniel & Cooke 14760 (18e). 1990 SIMSIA 85

McDonald 1010 (4). Padilla, M. 13 (17). McGregor, R. 16325 (14); 16728 (12); 16788 Painter et al. 14450 (12). (12). Palmer, Ed. 96 (8); 250 (14); 422 (17); 427 (17); McGregor et al. 821 (14). 440 (17); 471 (17); 493 (17); 602 (11); 610 McVaugh, R. 10585 (12); 20627 (17); 24401 (12); 611 (12); 612 (12); 613 (12); 615 (12); (17). 622 (17); 643 (12); 741 (10); 1005 (13). McVaugh & Koelz 10 (17); 76 (17); 104 (17). Palmer, E. J. 9790 (12); 12119 (12); 14539 (12); Mearns, E. A. 873 (13); 1232 (12); 1457 (12). 26793 (12); 33567 (12). Mears, J. 529E (12). Panero & G6mez 1099 (3). Mears & Mears 1490 (12); 3451 (14). Panero & Schilling 433 (7b); 448 (17). Medellin, F. 190 (17). Paray, C. C., & Palmer 471 (17); 472 (17). Meebold, A. 28030 (12). Parks et al. 52 (12); 173 (12). Melchert et al. 6259A (17); 6337 (17); 6428 (7b). Parry, L. 435 (17); 1307 (18a); 2193 (13); 2235 Messer, E. 72/lSla (13); 72/162 (13). (11); 2336 (8); 2565 (12); 2881 (14); 3023 Mexia, Y. 1832 (7a). (13). Miranda, F. 910 (18a); 1335 (13); 1572 (11); Parry et al. 560 (12); 561 (13). 2301 (18a); 2368 (18a); 2504 (13); 4808 Paul, Bro. 595 (18e). (18a); 5930 (18a). Paul 1 (12). Miranda & Barkley 16M978 (17). Peebles et al. 3377 (13). Moldenke, H. 7001 (12). Pennell, F. W. 5855 (3). Molina, A. 1413 (4); 13517 (11); 15512 (1); Pennington, C. 633 (17). 21030 (1); 21058 (17); 21326 (11); 23020 Perez, E. 435 (3). (4); 23027 (4); 23122 (4); 23157 (18d); Perez, G. 92 (12). 23238 (18d); 27306 (18d). Perez & Cuatrecasas 5874 (3). Molina & Molina 24566 (18d); 24819 (17); 24887 Pineda, A. 996 (17). (11); 24895 (1); 24913 (1); 24027 (17); Pinkava et al. 15537 (12). 24095 (17); 25110 (18d); 25217 (18d); 26302 Piper, C. V. 5427 (18e). (17); 26347 (17); 26505 (17); 26556 (11); Pittier, H. 4847 (18e). 26872 (11); 26938 (17); 27021 (11); 27799 Pohl, R. 4627 (12). (11). Porter, D. M. 1275 (18d). Molina & Montalvo 21575 (18a). Powell, A. M. 3445 (12). Molina et al. 16001 (11); 16163 (17); 30136 (11); Powell & Edmondson 534 (12); 599 (17); 603 30280 (17). (17); 640 (17); 808 (17); 840 (17); 998 (17). Moore, H. E. 1429 (17). Powell & Stroeder 83 (12). Moore & Wood 4662 (11). Powell et al. 1421 (12). Morales, J. 850 (18a); 1168 (18a); 1170 (11). Pringle, C. G. 321 (13); 1083 (13); 1738 (11); Moran, Y. M. 6 (17). 4044 (11); 4622 (17); 5090 (17); 8739 (12); Moreno, P. P. 18360 (18d); 18387 (18d); 22322 9897 (17); 9898 (18a); 11513 (11). (18d). Proctor, G. R. 21945 (18b); 24496 (18b); 32765 Moreno & Stevens 22882 (18d). (18b); 38472 (18b). Motte, E. 349 (11). Purpus, C. A. 1541 (17); 1542 (17); 3022 (11); Mueller, C. H., & Mueller 261 (12); 409 (12); 3226 (13); 3827 (17); 7472 (15); 9120 (11); 533 (12); 682 (12); 990 (12). 9121 (11); 9122 (11); 11121 (15); 13085 Mueller, F. 143 (13); 240 (13); 597 (13). (15); 13095 (15); 15022 (15); 16310 (11). Nealley, G. 95A (12). Pruski, J. 1269 (18b). Nee, M. 161 (17); 22203 (14); 22837 (17); 23485 Ramia 3027 (3). (17). Ramos, C. H. 266 (17). Nelson, E. W. 1426 (11); 1445 (11); 1937 (13); Ramos, F. M. 1345 (17); 77023 (17). 6603 (12). Ramos, F., & Romero 5 (17). Nuinez, J. 6660 (11). Raven & Gregory 12903 (12); 19248 (12); 19266 Oberwinkler & Oberwinkler 14120 (3). (12). Olsen & Escobar 496 (3); 497 (3); 600 (3). Read, J. A94 (12); 208 (12). Olsen & Lane 384 (18a). Reed, E. 596 (3). Opler, P. 574 (2). Reeves & Pinkava P13016 (12). Orcutt, C. R. 3947 (17); 4114 (17); 4334 (17); Reiche, K. 344 (18a). 5684 (17). Reichenbach & Smith 466 (18a). Ortega 25 (18c). Renson, C. 46 (18d). 86 SYSTEMATICBOTANYMONOGRAPHS VOLUME30

Reveal et al. 4171 (11). 7430 (12); 8405 (12); 16830 (12); 16989 Reverchon, J. 308 (12); 508 (12). (12); 17701 (12); 17707 (12); 17747 (12); Rincon, C. 8 (3). 19574 (12); 19630 (12); 31290 (12); 31753 Robert & Passini 5121 (12). (12); 31770 (12); 32669 (12); 32904 (12); Roberts & Keil 10456 (12). 32924 (12). Rock, H. M497 (12). Shreve, F. 8380 (12); 8827 (13); 8871 (13). Rodriguez, F. M. 1442 (16); 37856 (16). Skutch, A. 130 (17); 662 (17); 1143 (11). Rodriguez, J. 64 (12). Small & Wherry 11849 (12). Rogers et al. 6786 (12). Smith, C. L. 240 (11); 817. (11). Roman, E. 63 (17). Smith, H. H. 594 (3). Rosas, M. R. 467 (17). Smith, J. D. 2408 (18a). Rose, J. N. 2875 (13). Smith, L. C. 263 (18a); 854 (13); 909 (17). Rose & Hay 5808 (17). Smyth, E. G. 206 (17). Rose & Painter 6647 (17); 6816 (17). Socorro & Ferreira 2573 (16). Rose & Russell 24326 (12). Sodiro 34/2 (3). Rossbach, G. 4770 (12). Soejarto, D. D. 932 (3). Roybal, J. 746 (12); 803 (12); 989 (12); 1091 Solano, C. 384 (18a). (12). Solano & Vara 115 (11). Runyon, R. 302 (12); 349 (12); 1368 (17); 3355 Solis, I. 81 (12); 129 (12); 415 (17); 14864 (3); (12); 5991 (12); 6085 (12). 16401 (3). Rusby & Pennell 462 (3). Soto, J. C. 3474 (18a). Ryensky, D. 107 (17); 115 (17). Soto & Silva 1842 (18a). Rzedowski, J. 1514 (17); 1611 (17); 6828 (13); Soto et al. 8088 (12). 7440 (14); 11523 (17); 15830 (17); 16253 Spellenberg & Spellenberg 38311 (13). (18a); 16287 (17); 18109 (7a); 23516 (18a); Spooner, D. M. 2481 (14); 2482 (12); 2761 (11); 25475 (13); 29500 (18a); 29801 (11); 34353 2762 (17); 2763 (17); 2766 (11); 2767 (11); (17). 2768 (18a); 2769 (18a); 2770 (18a); 2772 Rzedowski & McVaugh 272 (7a). (11); 2780 (17); 2781 (1); 2785 (17); 2786 Safford, W. E. 1394 (17). (18a); 2787 (18a); 2788 (7b); 2789 (9); 2791 Sagastegui & Cabanillas 8740 (16). (11); 2792 (18a); 2794 (17); 2799 (14); 2800 Salvin & Godman 133 (1). (14); 2801 (14); 2802 (15); 2803 (15); 2804 Sanchez 2368 (16). (15); 2805 (15); 2806 (15); 2807a (15); Sanchez et al. 8352 (12). 2807b (15); 2808 (15); 2809 (15); 2811 (15); Sanders & Harborne 76054 (12). 2812 (14); 2814 (18c); 2815 (18c); 2816 (9); Sanderson, S. C. 198 (12). 2817 (13); 2818 (18a); 2819 (11); 2821 (13); Sandino, J. C. 266 (18d). 2822 (18a); 2824 (11); 2827 (11); 2829 Sauer, J. D. 1582 (17). (18a); 2831 (13); 2836b (13); 2880 (15); Saunders et al. 1452 (11). 2881 (15); 2882 (14); 2884 (14); 2885 (14); Schaffner, J. G. 19 (17); 162 (17); 265 (17); 389a 2886 (14); 2887 (14); 2900 (2); 2901 (4); (17). 2902 (18e); 2903 (18e); 2904 (18e); 2905 Schmitz 48 (17); 49 (17); 79 (17). (18e); 2906 (18e); 2907 (16); 2908 (16); Schott, A. 911 (15). 2909 (3); 2916 (3); 2918 (3); 2975 (3). Schreiter 1256 (16). Spooner & Brandenburg 2181 (12); 2188 (17); Schumann, W. 5 (17); 357 (17). 2189 (17); 2193 (17); 2194 (17); 2195 (17); Scudday, J. 551 (12). 2204 (17); 2209 (14); 2223 (14); 2229 (17); Seaton, H. E. 329 (17). 2230 (17); 2232 (17); 2247 (17); 2264 (18a); Seigler & Becker DS3088 (12); DS3344 (13). 2271 (17); 2274 (17); 2279 (7b); 2280 (17); Seigler & Lockwood DS8995 (12). 2292 (17); 2300 (18a); 2301 (18a); 2302 Seigler et al. DS8540 (12); DS2672 (12). (17); 2304 (17); 2305 (17); 2306 (17); 2311 Seler & Seler 584 (17); 684 (14); 1055 (12); 1185 (17); 2313 (17); 2324 (17); 2326 (17); 2336 (17); 1442 (11); 2178 (18a); 3027 (11); 3098 (13). (11); 4345 (13); 4472 (7a). Spooner & Burgos 2600 (13); 2601 (13); 2602 Sesse et al. 2867 (17); 2869 (13); 2870 (17); 2937 (13); 2603 (13); 2604 (11); 2606 (13); 2607 (18d); 2938 (17); 2944 (17); 2947 (11); 4021 (11); 2609 (13); 2615 (11); 2622 (13). (13). Spooner & Clausen 4584 (16); 4604 (16); 4608 Sharp, A. J. 441006 (17); 441223 (17). (16). Shiller, I. 847 (12). Spooner & Dorado 2627 (13); 2631 (13); 2635 Shinners, L. 7315 (12); 7363 (12); 7376 (12); (13x18a); 2636 (13); 2637 (18a); 2638 (13); 1990 SIMSIA 87

2639 (13x18a); 2641a (11); 2644 (13x18a); (6); 43108 (1); 47294 (11); 51470 (11); 2645 (13x18a); 2646 (13); 2649b (11); 2654 51625 (11); 52182 (18d); 57062 (3); 62345 (17); 2656 (13); 2658b (11); 2659 (13); 2664 (3); 62424 (3). (18a); 2666 (18a); 2667 (18c); 2669 (18c); Strother, J. 97 (12); 268 (12); 860 (12). 2670 (17); 2673 (17); 2674 (1); 2675 (1); Stuckert, T. 6403 (16). 2681 (11); 2683 (1); 2684 (17); 2684b (11); Stuessy, T. 909 (12). 2687 (17); 2688 (18a); 2690 (5); 2691 (18a); Stuessy & Gardner 3008 (18a); 3108 (7a); 3145 2691a (18a); 2692 (5); 2693 (18a); 2695 (13); 3151 (11); 4039 (14); 4041 (14); 4083 (18d); 2696 (18d); 2697 (18d); 2699 (18d); (17); 4095 (17); 4111 (18a); 4192 (17); 4196 2700 (18d); 2701 (4); 2702 (11); 2703 (18d); (17); 4211 (13); 4225 (7a); 4239 (13); 4259 2704 (4); 2714 (4); 2723 (18d); 2740 (18d); (17); 4261 (18a); 4262 (13); 4271 (11); 4304 2746 (5); 2747 (18a); 2748 (6); 2749 (18a). (17); 4314 (18a); 4316 (11); 4335 (17); 4345 Spooner & Guevara 2921 (3); 2923 (3); 2924 (3); (1); 4377 (18a); 4552 (18d); 4561 (18d); 2925 (3); 2927 (3); 2933 (3); 2935 (3); 2936 4573 (18a); 4578 (17x18a). (3); 2937 (3); 2938 (3); 2941 (3); 2944 (3); Stuessy & Meacham 3511 (12). 2945 (3); 2952 (3); 2953 (3); 2956 (3); 2957 Stuessy & Roberts 3635 (14). (3); 2972 (3); 2973 (3). Sundberg & Lavin 2759 (17); 2922 (17); 3009 Spooner & Linder 2750 (5); 2753 (1); 2758 (13); 3020 (11); 3067 (11). (18a); 2759 (5). Sundberg et al. 1744 (17); 2404 (18a); 2417 Spooner & Schilling 2453 (17); 2455 (17); 2456 (18a). (13); 2457 (13); 2458 (13); 2459 (13); 2461 Surratt, H. 110 (12). (13); 2463 (17). Templeton, B. C. 7576 (18a). Spooner & Triplehorn 1058 (12); 2047 (17); 2048 Tharp, B. C. 43-901 (12); 43-902 (12); 43-903 (17); 2073 (17); 2098b (13); 2100 (13). (12); 43-904 (12); 1221 (12); 1881 (12); Spooner & Villasenior 2488 (17); 2489 (13); 2490 3831 (12); 8834 (12). (17); 2491 (13x 17); 2494 (18a); 2495 (11); Tharp & York 49271 (12). 2504 (18a); 2505 (11); 2528 (10); 2535 Tharp et al. 51-1647 (12). (17x18a); 2537 (11); 2538 (17x18a); 2546 Thurber, G. 953 (13). (17); 2550 (8); 2557 (17); 2562 (7b); 2563 Tidwell, C. 111 (12). (7b); 2565 (7b); 2566 (7b); 2567 (11); 2568 Tolsted, W. 7249 (12). (11); 2569 (18a); 2570 (11); 2571 (17); 2672 Ton, A. S. 838 (17); 1070 (17); 1231 (17); 1304 (11). (17); 1562 (17); 2033 (18a); 2060 (11). Standley, P. C. 14009 (4); 19483 (18d); 19447 Tonduz, A. 508 (13). (18d); 21639 (18d); 26385 (18e); 27148 Torres, A. M. 1526 (12). (18e); 29209 (18e); 29788 (18e); 32098 Traverse, A. 1788 (12). (18e); 58037 (13); 58575 (11); 59379 (1); Triana, J. 1361 (3); 1362 (3). 60288 (11); 60374 (13); 60411 (18a); 60675 Troll, C. 777 (17). (18a); 61158 (1); 61199 (1); 61469 (18a); Trujillo, B. 6356 (1). 62389 (17); 62834 (11); 62986 (11); 65054 Turner, B. L. 37 (17); 80A-25 (11); 477 (12); (1); 66066 (17); 66164 (17); 75023 (11); 4556 (12); 4583 (12); 4953 (12); 9232 (12); 75118 (18d); 75585 (11); 76514 (18a); 77582 15196 (17); 15197 (17); 15198 (17); 15212 (18a); 77667 (18a); 77857 (18d); 79977 (11); (17); 15265 (17); 15415B (17). 80307 (17); 80861 (11); 81339 (11); 81928 Turner & Barrie 15336 (17). (11); 81996 (17); 83164 (17); 83944 (17); Turner & Powell 1046 (12); 1093 (14). 85789A (17). Turner & Tapia 15522 (15). Standley & Padilla 3179 (18d); 3463 (18d); 3549 Turner & Turner 80-26M (14). (18d). Tyson, E. L. 2697 (18e). Standford et al. 2287 (12). Tyson & Blum 2595 (18e). Steere, W. C. 1026 (15); 1625 (15); 3027 (15). Tyson et al. 3024 (18e); 3069 (18e). Steiger, T. L. 938 (13). Ugent, D. 1396 (12). Steinbach, R. 711 (16). Ugent et al. 1381 (17); 2630 (13). Stevens, W. D. 4768 (18d); 5744 (18d); 10921 Urbatsch, L. E. 928 (12); 935 (12); 937 (12); 965 (18d); 18997 (18d). (12); 2407 (12); 2412 (12); 2417 (12); 2804 Stevens & Grijalva 15708 (18d). (13). Stewart, R. 1373 (13); 1481 (12); 1643 (12); 1706 Urbatsch & Pridgeon 3014 (11). (12). Valentin, N. 162 (17). Steyermark, J. A. 30968 (18d); 32394 (17); Vasquez, J. 4988 (18a). 32894 (17); 32982 (11); 36806 (17); 42931 Vaughan, J. 512 (12). 88 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 30

Vaughan et al. 1154 (14). Weberbauer, A. 5309 (16); 5934 (16). Vega, R. 654 (17). Webster, G. 506 (12); 16984 (11). Ventura, A. 150 (17); 1062 (17); 1154 (17); 1308 Webster & Armbruster 21031 (18c). (17); 1723 (17); 2531 (11); 4110 (17); 4121 Webster & Breckon 15472 (14); 16132 (17); (11); 8709 (11). 16216 (13). Ventura, E. 1414 (17). Webster et al. 17M553 (13); 11173 (12). Venturi, S. 4166 (16); 4999 (16); 6277 (16). Weedon, R. R. 4021 (12); 4101 (12). Vergara et al. 8578 (12). West, R. C. J23 (17). Vibrans, H. 157 (17); 410 (17); 413 (17); 516 Whiffin & Rodriguez 206 (14). (17); 608 (17); 674 (17); 718 (17); 840 (17). White, S. S. 3120 (10); 3292 (10); 4155 (10). Viereck, H. W. 330 (12). White & Chatters 106 (12). Villarreal, L. M. 196 (13); 1264 (18a); 1479 (11); Whitehouse, E. 7059 (12); 10102 (12); 10110 2133 (18a); 6997 (17); 7027 (18a). (12); 11026 (12); 11027 (12); 11031 (12); Villasefior, J. 54 (17). 16717 (12); 16718 (12). Villasefior & Spooner 679 (18a); 684 (11); 723 Wieder et al. 6 (12). (11); 742 (8); 762 (7b); 766 (7a); 767 (11); Wilkinson, E. 41 (12). 772 (18a). Williams, R. S. 242 (18e). Villasenior et al. 269 (11); 292 (8). Williams & Molina 15188 (18d); 15410 (11); Villegas, M. D. 3 (17); 22 (17); 47 (17); 150 42327 (4); 42449 (18d). (17); 174 (17); 187 (17); 229 (17); 253 (17); Williams et al. 21881 (11); 22275 (11); 22321 274 (17); 608 (17); 707 (17); 782 (17); 902 (17); 22535 (11); 25231 (1); 26125 (17); (17). 26135 (17); 26142 (1); 26916 (17); 27275 Voorhies & Sanchez 15-12 (18a). (11); 41109 (18a); 41160 (11); 41210 (1); Wallace et al. 131 (11). 41276 (11); 41366 (18a); 41437 (17); 42168 Warnock, B. W264 (12); 5946 (12); 6015 (12); (1); 43367 (1). 6187 (12); 6718 (12); 6727 (12); 6767 (12); Williges, G. G. 366 (12); 416 (12). 6818 (12); 8887 (12); 9222 (12); 11219 (12); Wilson, E., & Johnston 8591 (17); 8592 (17). 11292A (12); 11363 (12); 11372 (12); 12562 Wilson, J. S. 12184 (12); 12429 (14); 12638 (12); 13436 (12); 14541 (12); 14758 (12); (12). 14893 (12); 15223 (12); 15764 (12); 19467 Wolff, S. 3273 (12). (12); 20202 (12); 22144 (12); 22239 (12); Woronow, G. 884 (18a); 2047 (17); 2804 (17); 23531 (12); 46523 (12); 47137 (12). 2433 (18a); 2576 (18a); 2584 (18a). Warnock, B., & Cameron 9879 (12). Worth, M. G. 23 (14). Warnock, B., & Johnston 17132 (12). Wright, C. 204 (12); 330 (12); 331 (13); 1224 Warnock, B., & McBryde 14541 (12); 14758 (13). (12); 14893 (12). Wunderlin, R. P. 5081 (18b). Warnock & McVaugh 47182 (12); 47237 (12). Wussow & Landry 331 (13). Warnock, M. J. 2564 (11). Wynd & Mueller 177 (12). Waterfall, U. T. 5439 (12); 16660 (12). York, H. 404 (12). Waterfall & Wallis 14300 (14). Zarco, J. 12 (17). Watson, T. J. 540 (12). Zer6n & Villasenior 283 (17). Wawra 1088 (11). Zizumbo, D. Z723 (13). Weaver, J. N. 883 (17). Zizumbo & Colunga 288 (18c). Weber, R. 245 (17); 290 (13); 295 (13); 300 Zola, M. G. 12 (15). (13). Zuchowski, W. 1-80a (17).

INDEX TO SCIENTIFIC NAMES

Accepted names are in roman type; themain entry for each is in boldface. Synonyms are in italics.

Armania Bertero ex DC. 23 Axiniphyllum Bentham 17 fruticulosa (Sprengel) Bertero ex DC. 23, Barrattia A. Gray & Engelmann ex A. Gray 29 21, 23 Aspilia calva A. Gray & Engelmann ex A. Gray grosseserrataM. E. Jones 45, 49 23, 49 1990 SIMSIA 89

Coreopsidinae 3, 17 Milleria L. 17 Coreopsis Montanoa Cerv. 20 amplexicaulis Cavanilles 60, 64 Phoebanthus S. F. Blake 3 foetida Cavanilles 2, 64, 68 Salmea heterophylla Cavanilles 73 fruticulosa (Sprengel) Steudel 29 Cosmos Senecio L. 6 hintonii Sherff 45 Sigesbeckia L. 17 Echinacea Simsia R. Brown 23 heterophylla (Cavanilles) D. Don 73 Simsia Persoon 23-24 Encelia Adanson 1, 3 sect. Geraea (Torrey & A. Gray ex A. Gray) sect. Simsia (Persoon) A. Gray 23 A. Gray 73 amplexicaulis (Cavanilles) Hemsley 60 abrotanoides (Meisner) F. v. Mueller 73 adenophora Greenman 65 adenophora (Greenman) S. F. Blake 65, 68 calva (A. Gray & Engelmann ex A. Gray) A. amplexicaulis (Cavanilles) Persoon 1, 2, 5, 11, Gray 49 16, 17, 19, 20, 21, 23, 24, 49, 55, 60-64, chaseaeMillspaugh 57 65, 67, 68, 78, 79, 80, 81 cordata (H. B. K.) Hemsley 61 var. decipiens (S. F. Blake) S. F. Blake 61, deltoidea Rusby 31, 34 64 exaristata (A. Gray) A. Gray ex Hemsley 52 var. genuina S. F. Blake 60 foetida (Cavanilles) Hemsley 64 anethifolia R. Brown 73 frutescens (A. Gray) A. Gray 73 annectens S. F. Blake 5, 17, 19, 22, 23, 37-40 fruticulosa (Sprengel) Hieronymus 30 var. annectens 2, 5, 20, 21, 23, 24, 38-39, ghiesbreghtii A. Gray 26 79, 80, 81 grandiflora (Bentham) Hemsley 70 var. grayi (Sch.-Bip. ex S. F. Blake) D. M. heterophylla (H. B. K.) Hemsley 61 Spooner 25, 38, 39-40, 79, 80, 81 hirsuta Kuntze 59 auriculataDC. 61 f. radiataKuntze 59, 60 calva (A. Gray & Engelmann ex A. Gray) A. hispida (H. B. K.) Hemsley 73 Gray 5, 6, 11, 16, 19, 20, 21, 23, 24, 44, lagascaeformis (DC.) A. Gray ex Hemsley 52 49-52, 79, 80, 81 mexicana Martius ex DC. 61 var. subaristata (A. Gray) S. F. Blake 49 pilosa Greenman 52 canescens (Torrey & A. Gray ex A. Gray) A. polycephala (Bentham) Hemsley 71 Gray 73 purpurea Rose 52, 55 caucana Cuatrecasas 31 sanguinea (A. Gray) Hemsley 45 chaseae (Millspaugh) S. F. Blake 2, 5, 6, 7, 9, var. palmeri A. Gray in S. Watson 45 10, 11, 12, 13, 14, 15, 16, 17, 18, 19, 21, scaposa (A. Gray) A. Gray 73 22, 24, 25, 53, 57-58, 60, 65, 79, 80, 81 sericeaHemsley 26 cordata (H. B. K.) Cassini 61 sodiroi Hieronymus 31 cronquistii H. Robinson & R. D. Brettell 69 subaristata (A. Gray) A. Gray ex Hemsley 49 dombeyana DC. 5, 6, 7, 9, 10, 11, 12, 13, 14, tenuis Fernald 40 17, 18, 19, 20, 21, 22, 24, 32, 58, 59-60, Enceliopsis A. Nelson 3 65, 79, 81 Flourensia DC. 3 erythranthemaSch.-Bip. ex A. Gray 45 Geraea Torrey & A. Gray ex A. Gray 3, 73 eurylepis S. F. Blake 1, 2, 5, 6, 9, 10, 11, 13, canescens Torrey & A. Gray ex A. Gray 73 14, 15, 16, 19, 20, 21, 22, 23, 24, 25, 53, Heliantheae 3 54, 56-57, 58, 79, 80, 81 Helianthus L. 3, 4, 6 exaristata A. Gray 19, 21, 52, 54, 55 amplexicaulis DC. 61 var. epapposa S. F. Blake 52 hastatus Sesse & Mocinlo 61 var. perplexa S. F. Blake 52, 55 trilobatusLink ficifolia Persoon 64 sericeus Sesse & Mocifio 61 foetida (Cavanilles) S. F. Blake 1, 2, 3, 5, 6, Hopkirkia 7, 8, 9, 10, 11, 13, 14, 15, 16, 17, 19, 20, fruticulosa Sprengel 29 21, 22, 23, 38, 39, 45, 57, 58, 60, 64-73 Iostephane var. decipiens S. F. Blake 61 heterophylla (Cavanilles) Bentham ex Hemsley var. foetida 2, 8, 9, 10, 11, 14, 15, 16, 18, 3, 73 19, 20, 22, 25, 64, 65-68, 69, 70, 72, 79, Liabeae 17 80, 81 90 SYSTEMATICBOTANYMONOGRAPHS VOLUME 30

var. grandiflora (Bentham) D. M. Spooner schaffneri Sch.-Bip. ex A. Gray 61 4, 8, 9, 13, 16, 19, 20, 25, 67, 68, 69, 72 sericea (Hemsley) S. F. Blake 21, 26, 28 73, 79, 80, 81 setosa S. F. Blake 2, 5, 11, 13, 16, 17, 21, 22, var. jamaicensis (S. F. Blake) D. M. 26, 42-45, 79, 80, 81 Spooner 2, 8, 9, 11, 19, 21, 25, 65, 67, simplex (Lindley) F. v. Mueller 74 68-69, 81 sodiroi (Hieronymus) S. F. Blake 31 var. megacephala (Sch.-Bip. ex S. F. Blake) steyermarkii H. Robinson & R. D. Brettell 1, D. M. Spooner 8, 9, 11, 17, 19, 20, 21, 5, 16, 19, 21, 22, 26, 27, 35, 57, 80, 81 25, 67, 69-70, 79, 80, 81 subaristataA. Gray 21, 49, 52 var. panamensis (H. Robinson & R. D. submollicoma S. F. Blake 56, 57 Brettell) D. M. Spooner 2, 8, 9, 11, 19, subsetosaH. Robinson & R. D. Brettell 38, 39 20, 21, 24, 32, 67, 71, 72-73, 79, 81 tenuifoliaR. Brown 74 frutescensA. Gray 73 tenuis (Fernald) S. F. Blake 2, 5, 16, 19, 25, fruticulosa (Sprengel) S. F. Blake 5, 11, 16, 39, 40-41, 80, 81 17, 19, 20, 23, 25, 28, 29-34, 35, 79, 81 teretifolia (Meisner) F. v. Mueller 74 ghiesbreghtii (A. Gray) S. F. Blake 5, 11, 21, triloba S. F. Blake 45 22, 24, 25, 26-28, 33, 79, 81 villasenorii D. M. Spooner 2, 5, 13, 17, 19, grandiflora Bentham 68, 70 20, 22, 25, 29, 39, 41-42, 43, 80, 81 grayi Sch.-Bip. ex S. F. Blake 39 Stirlingia grisea S. F. Blake 31 abrotanoidesMeisner 73 guatemalensis H. Robinson & R. D. Brettell latifolia (R. Brown) Steudel 73 66 simplex Lindley 74 heterophylla (Cavanilles) Persoon 61, 73 tenuifolia (R. Brown) Steudel 74 heterophylla (H. B. K.) DC. 2, 61, 73 teretifoliaMeisner 74 hintonii H. Robinson & R. D. Brettell 38, 39 Tithonia Desf. ex Gmelin 3, 17 hirsuta (Kuntze) S. F. Blake 19, 59 Verbesina L. 3 hispida (H. B. K.) Cassini 73 argentea Bertoloni 26 holwayi S. F. Blake 1, 2, 5, 25, 27, 35-36, 54, argenteaGaudichaud 26 79, 81 Viguiera Kunth 2, 3, 4, 6, 18, 21, 22, 28, 78 jamaicensis S. F. Blake 69 ser. Dentatae S. F. Blake 4 kunthiana Cassini 61 ser. Grammatoglossae S. F. Blake 3, 4, 5, 6, lagascaeformis DC. 1, 2, 5, 11, 13, 15, 16, 17, 21 19, 20, 21, 22, 23, 24, 25, 35, 52-56, 57, ser. Viguiera 4 67, 68, 79, 80, 81 cordifolia A. Gray 4, 18, 80 latifolia R. Brown 74 grammatoglossa DC. 4 megacephala Sch.-Bip. ex S. F. Blake 69, 70 mima S. F. Blake 4 molinae H. Robinson & R. D. Brettell 2, 5, ovata (A. Gray) S. F. Blake 4, 21, 28 11, 20, 26, 27, 33, 34-35, 79, 81 parkinsonii (Hemsley) S. F. Blake 4 panamensis H. Robinson & R. D. Brettell 72 potosina S. F. Blake 4 pastoensis Triana 30, 33 pringlei Robinson & Greenman 4 polycephala Bentham 71 seemannii Sch.-Bip. 4 pubescens Triana 30 sessilifolia DC. 74 sanguinea A. Gray 5, 6, 13, 16, 17, 20, 21, 22, Ximenesia Cavanilles 3 23, 24, 25, 45-49, 79, 80, 81 cavanillesii Sprengel 73 subsp. albida S. F. Blake 45, 49 cordata H. B. K. 61 var. palmeri (A. Gray) S. F. Blake 45 foetida (Cavanilles) Sprengel 64 santarosensis D. M. Spooner 2, 11, 17, 19, 25, heterophylla H. B. K. 61 27, 29, 30, 42, 79, 81 hirta Martius ex DC. 61 scaposa A. Gray 73 hispida H. B. K. 73